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GENE? CS AND CYTOLOGY OF SOLE InPORTANT

mmuuoor True mam 1x§£UfIVE OF T1412 UIIITSD S'l‘ATIflS

by

Scott Eguley

A ThES 8

Submitted to the Graduate School of Richigan

State College of Agriculture and Applied

Science in partial fulfilment of the

requirements for the degree of

1‘.‘;A13'l‘ER OF SC MICE

Department of BO”A?Y

1942

ILTRODUCTIOH

Foresters during recent years have become increasingly

aware that the forest practices of 1900 are in some instan-

ces no longer sound. The changing concepts are evidence of

rational and normal develOpment for they have paralleled the

rapid accumulation of facts collected by numerous workers in

the related bioloaical and physical sciences.

Of all concepts and techniques which have been altered

or revised none is more important than the changing attitude

with regard to the fundamental unit with which the forester

works. Gradually the concept that the species is the unit

of forest practice is being destroyed. The revolutionary

work in genetics of the past forfiy years has demonstrated

beyond any reasonable question of doubt that foresters, no

more than growers of annual crOps, can any longer well af-

ford to ignore the genetically superior members of a species.

Although the study of the genetics and cytogenetics of

forest trees is of comparatively recent origin, an extensive

and valuable literature has develOped. In addition, the

older as well as contemporary work in related fields con-

tains much valuable information and technique which is of

value to the forest tree breeder.

This paper represents a preliminary effort to collect

and evaluate in some degree of unity the contributions of

numerous workers to the end that their inferences and con-

clusions may be made more readili available to the genetic-

..4 45‘“); '.3

Jink \5 a t

-11-

ist and cytologist interested in the improvement of our for-

est trees. The critical review, as a means of systematizing,

evaluating, and preserving the rapidly accumulating data in

all fields of science, is receiving reCOgnition as a highly

desirable research technique.1

The genetic and cytogenetic aspects of eight of our im-

portant har‘wood forest tree genera have been treated in this

review. Each genus has been discussed as a unit with empha-

sis upon the following principle tOpics: (l) artificial se-

lection from wild stock; (2) controlled breeding and selec-

tion; and (3) cytology. The "literature cited” for each

genus follows the discussion of that genus. In addition a

list of "literature not cited" has also been indicated. In

the latter list are noted publications which were not avail-

able to the author or they represent pOpular articles review-

ing scienctific papers which have been consulted. This

arrangement has been adopted for the purpose of broadening

the scope and usefulness of the review.

Sec: Bradford, F. C. (1942) Conserving our harvests.

Science 95 (2471): 465-469.

-iii-

CONTENTS

Page

Introduction .................................. ii

ForULUS ....................................... 1

Artificial selection from wild stock ........ 4

Controlled breeding and selection............ 8

Cytology .................................... 52

Literature cited ............................ 55

SI'XLIX 000......IOIOOOQOOOOQOOOOOOOO00......O... 6]—


Controlled breeding and selection ........... 65

CEItOJ-OsrIO...0....OOOOOOOOOOOO.00.00.0000... 69


JUGLANS ....................................... 79

Artificial selection from wild stock ........ SO

Controlled breeding and selection ........... S2

Cytology .................................... 88

Literature cited ............................ 9O

CARYA ......................................... 95



Cytology .................................... 97


BETULA ........................................ 101



Cytology .................................... 107


QUERCUS coco-ooooooooooooooococo-0.000.000.0000 115



Cytology .................................... 125


UIAUS ......................................... 129



Cytology .................................... 156


ACER .......................................... 141



Cytology .................................... 148


-iv-

SALICACEAE Lindley

Killow or Peplar Family

The Salicaceae, with more than 200 species, includes 5

1, 1genera -, Populus L., Salix L., and 0110967118. I‘Iaka

Ho

. The lat-

ter, a monotypic genus, is restricted to northeastern Asia

(Rehder, 1940).

Salix and Ponulgs have been reported in the upper Cre-

taceous of North America, Greenland, western EurOpe, Japan,

and Russia and the Cenozoic history of the family is rich in

variety (Larrah, 1959). They are at present most abundant in

the temperate regions of the northern hemiSphere.

Although a timber contributing family of minor importance,

a number of species are valued members of pioneer types which

play significant roles in soil and water conservation. Since

many species are easily reproduced vegetatively and produce a

network of fast-growing roots they are used for erosion con-

trol. Some of the distinctive forms are widely used as orna-

mentals (Harlow a harrar, 1941).

POPULUS L. (pOplar)

POpulus L., represented by about 50 tree species is, for

the most part, confined to the temperate regions of the north-

ern hemisphere. Fifteen species are native to North America

1 According to harlow a harrar (1941), five genera are now

recoanized due to the recent work of Japanese taxonomists.

Toisusu, previously included under Salix, has been raised

to generic rank. The genus Turanra_EEE—been added. Both

are exclusively Asiatic in d-s ribution.

but only about five1 of these occur in commercial size and

quantity in the United States (Harlow L harrar, 1941).

Tte 3enus is diVided into five e etiors: (1) Lance Duby

(white pOplars, aspens); (2) Leucoides Spach; (5) Tacamahaca

Spach (balsam poplars); (4) Aeneiros Luby (cottonwoods, black

neplars); and (5) Turanra Bun3e (Petder, 1940).

The wood of the American and EurOpean species is employed

in considerable quantities for pulp, excelSior, veneer, lumber

for boxes and various other small articles of woodenware (Brown

& Panshin, 1940; Sargent, 1922). The high orn11ental value of

many of the species has resulted in their wide use as street

and shade trees. The white peplar (E. 3123 L.) and its several

varieties, the Chinese peplar (2. sinonii Carriere), and the

\4

lomhardy peplar (P. hi ra ita lica Luenchausen) have been ex-

tensively planted in the United States.

Unfortunate}.v considerable taxonomic difficulty has arisend

.‘

at various times amon3 systematic workers and forest tree bree‘-

ers With regard to the proper nomenclature to be used for cer-

tain species, varieties, forms or hybrids of the several poly-

morphic and easily hybridized species of this 3enus. A number

of workers have striven to bring some order out of this chaos.

piLn Great Uritrin Elwes and Henry (1915), henry (1910, 1914a

1914b, 1950) and Cansdale (1959) have contributed to taxonomic

clarification of the especially difficult and confused section

1P. trenilojdeg Licnaux, P. gparcideniaia fichaux, P. ta camahaea

Filler, P. trlsiocahpa 'nstata henry, var., P. delto ioes LarSuall.

Aejeiros Duby (cottonwoods and alach peplars). Houtzagers (1957)

in Holland, Quairiére (1956) in Bel3°um, and Sargent (1896,

192?) and dehder (1940) in the United States have also proviCed

valuable revisions and revaluations in nonenclature for the

whole genus. Loutzagers' work, especially, will prove an in-

valuable aid to breeders in ibis genus.I

In accord tith the olicv practiced in other sections of’3

this review, an attempt has been made in most cases to indicate

the probable identity of the many meaningless common or scien-

tific names (1ackin3 authority names) which have been used with

such promiscuity by some authors. rhe work of Rehder (1940)

has been used as reference and autLority on taxonomic questions.

It is also a matter of some taxonomic importance to rec03-

nize, as pointed out by Stout (1929) and Stout and Schreiner

(1955), that certain of the natural, or early described, hybrids

in this genus have been exclusively prepagated as clones.._,.

these clones have masqueraded as species undefi such names as

"P. serotina Hartig" (= X E. canadersis serotinc (Hartig) Fehder),

. T ‘ - o

VET. and "P. rc3enerata henry‘ (- X P. canadenSis regenerate

(Sc?.n1eider) Rehder, var.).

Most members of the genus are characterized by rapid

growth, ease of prepagation by twi3 or root cuttin3s, and winter

hardiness. All are wind pcllinatedl, essentially dioecious,

hybridize with case (at least within and between most sections),

and are capable of successfully maturing immediately viable

seed on cut twigs in the greenhouse. Many Species however5 ’

1The genus Salix L. is entomOphilous.

-5-

are hosts to a number of fungus diseases and insect pests.

With this general set of conditions it is not remarkable that

considerable interest has been directed to this genus by for-

est genet1c ists, both in tno United States and abroad.

is”:1’1':

ICIAL SELECTICI FRO; XILD STOCK

Considerable eiWioiShas been directed in EurOpe, especially

in France (Stout, 1950; Ackers, 1958), to the sele etion of na-

turally occurring hybrids, varieties, or forms of peplars supe-

rior for use in forest plantations. The business of growing

these trees in France as a timber crop to meet local demands

of wood using industries is claimed by these authors to be highly

successful.

In France as well as in Germany, however, according to

Cansdale (1938), the nomenclature in the genus is more than or-

dinarily confused. This condition, of course, has not materi-

ally advanced selection efLor ts or the ac chulation of reliable

data. Stout reported t‘ne three clones which are said to be the

most extensively used for forestry purposes in France as:

Eégénéré , "Angulata" and Peuplier a Ecorce Noire". we

probably 2 g. regenerata Henry 3 X 3. cnnwnrnis regenerate

(Schneider) Rehder but to suggest synonyms for the others re-

quires departure from the field of probability.2

Increasing interest in the possible use of peplars for

lRe;3nier (1934) has reportedly provided classification and de-

scription of many peplarsgrown in France as a preliminary step

in a breeding research program desilgned to isolate diseaase re-

sistant forms.

2Stout stated these names were best he was able to obtain from

French growers.

forestry purposes h5+s been manifest in inl_”nd in recent years.

Lotbiniere (l9?”5)and AC’ers (l 58) believe the growing of pep-

lars under forest management to be ncorowicoliv sound provided

de rable stom1 is used. Such ideas have encouraged search for

the most desirable wild forms best suited to the pla of manage-

ment and utilizati_on. ,

An initial and most fundamental step in that direction has

been accomplished by Cansdale (19 ’b) in his "”1 ey to t}.1e species

and hybrids of Poyulus cultivated in Britain". An assemblage

5:

of other 3 .nple.entary evaluating information on individual mem-a

bers of the genus has also been made by this author.

Cansdale and AcLers believe that X P. re enerata Henrv. _ w J

(3 X P. canadensis rege.erata (Schneider) Rehder) may well prove

to be a valuable timber producer in the British Isles as, sup-

posedly, it has in France. This clone has been grown in forest

plantations in Ensland for a full rotation (40 - 60 years) in

several localitie swith high yiield.

According to Ac :ers other native clones or introductions,

'1

including eugenii", ”robusta" and "3 I H "O .Y'N, I

generosa are pPOMlSlJb for

forestry use. The Species P. vunnanensis Dode of China and P.

trichocarpa Hooker of the Pacific coast region of North.America

are being tested in forest plantations on various sites in the

British Isles.

Interest in pOplar cultivation and breeding in Germany is

reportedby .'e1;tstein (19'65-a) and the resulting demand for

superior Wild stock for breedinak)

or direct use has greatly stim-

ulated wild selection.

Wettstein (1954) indicated that studies of climatic races

in various species of Poonlus and other 5enera have been carried

on by the German Association for the Breedin; of Forest Treesl.

The further refinement of progeny tests and seed certification

of isolated true breeding strains has also been instituted as

an aid to the work.

Isolation of at least two climatic strains of E. tremula L.

was later reported by Wettstein (1957-a) and reports of pr05eny

studies, showin5 variation amonr the seedlings of P. tremuloides“‘0

Michaux and P. tremula L. were made Ly the sane author (1957-0).

Vill (1930) reported that the hybrid X P. charkowiensis

Schroder (probably X P. canadens is Eu5enei (bixron-Louis ) Schelle)

is bein5 51rown extensively in the Fhineland. Althou5h the tree,

under plantation conditions, is said to make very 500d 5rowth

and to show considerable promise as a timber producer, it is,

nevertheless,subject to canker.

The discovery in Sweden of 9 clones of natural triploid

P. tremula L. (Nilsson-Ehle, 1936; Blomqvist, 1957; Tomator

'd

1957; Johnsnon, 1940) gives promise of supplying valuable Wild

material for direct forestry use in that country. According to

Nilsson—Ehle (1956, 958-b) the first triploid clone (discovered

near Lille, Skgne, South Sweden) grows more rapidly, produces

more wood per year, and is believed to be more resis ant to at-A

tack by Polyporns so. than the normal diploid European aspen.L

Also, because of its ”stronger growth", it is reported as able

to succeed in competition with lime, maple, elm and other hard-

woods. The native diploid on the other hand is said to be found

1Formed by Erwin Baur in 1932.

-5-

o 'I 1 '-. a ‘ .‘I +r

onlv on the outerr11.115in of such mixed 1-31.rd-.10od standS 15‘- 0014? ern

Swede.. Tests of the triploid or "515as" form have revealed its

value for use in the match and paper pulp industrie.. The form

is reputedly easy to reproduce by root cuttings and since a di-

-loid X trinloid cross has already croduced tetraoloid forms itp ‘ 4- L

is probable that triploid seed can be produced at will in the

future by diploid‘-

f"!

tet acloid crosses.1V4

b

In the United States the various wild species and hybrids

of Penulus, both native and exotic, are claimed by Stout and

Schr iner (1955) to be incapable of fully satisfyin5 the demands

for direct use in reforestation project. In Cznada, presumaoly,

the same situation is believed to exist. Peto (1958), however,

reported ,hat the natural hybrids found by Eeimburger (1956)

near Ottawa show considerable promise for use in forestry. Of

these natural c11os s<s, involvir5 the e::otic P. Elba L. and the

two native aspens (P. tremuloides hichaux and P. crandidentata

Michavx), the alba X 5randidentata hybrids are especially pro-

mising.

Stout and Schreir1er criticised the slow 5rowth and poor

rooting ability of the native P. tremuloides and P. crandidentata

and pointed out that many of the imported species and tytrids

are not hardy in the severe climate of the northeastern United

States. It is also claimed that althou5h some of the natural

hybrids, which are successfully 51oun in EurOpe, are 115hly de-

sirable from the standpoint of growth, reproducti-on, and physi-

cal characters of the wood they cannot survive the fun5us dis-

eases to which they are subjected in the new world (Schreiner,

-7-

Stout,

Artificial selectiow

been particularly fruitful

dantly demonstrated by the

wideely used as ornawentals

of the northern hemisphere.

CL]

With the

ments of Klotzscb, kegs,

of Van Fleet and

project was

Oxford Paper Company, Rumford,

Hew York Botanical Garden.

by Stout

(1957). R. H. ILCI’LQC,

ary report,

ject.

The preliminary 0bM3

evaluate and test the

the basis of their value for pulp\!ood re forestaiion

duce tbrou5h hybridiz

existence.

Stout and

th a t the c 01711:} on

from

for the

THU" 19 I" 017. S

.\| -. -L .3?) . 1.| \4.. I

;.Tr:T.(JJ‘l—JLD 12.1 '

exception of

)

Henry”,

started in

and Schreiner (lCR7, 1933, 1954-a,

a c 0- author

is credited with conception

active 0: tie

WHOM)“.

gation more valuable

Schreiner (1935), as

wild material in this genus has

hortimtuLz1risst as; s abunr

named varieties and forms

irl the: te'w1e1" te re .ic>nsJ

especially

‘1‘:Tv ‘ T: i '7‘an INK-

L "U U4._.Ja‘_4._j _L \.’i_'\-1

H L"

‘1‘

V"

the early tree hybridization eXpepi-

I

and the chestnut breedine work

Graves, the first comprehensive tree breeding

the 5enus Pepulus L. in 1924 by the

Raine in ollaboration with the

was conducted and pepOPtOd

1934b) and Schrei 1183’."

of the fir t (1927)C0

of tLe idea

breedin5 pro5ram was to

soecies and hybrids of Ponulus on.. U ‘

and to pro-

types than those in

previously noted, concluded

of the northeast (P. tremuloides and

a nd ca nk’e r-

lEspecially nelaflnsora medusae Thfim” a rust fun5us,

jlulthlLUV “ sa

nr1inatur1(lghotsohe) Che., :Juilflect'ii-vlli_

forming soecics

13331.

“Tel?" (1914) artificially

and P. anerlata auth9 3 P—Al‘_ —

Lybridized P. tricbocarba

1‘11 srvu~5d0 Iij;., E1anrflon‘1 Q3

5ena Ewes. (Boyce,

Hooker

erer’osii Henry.

i..)o

P. 91'15'1'r‘w‘iu‘1'e1t9t9) were too slow; :1 9.111132. too 110(1' in root-- -._-—. —.._.—_..

f

inn ability to be desirable f0“ reforest9tion 1M1ntin9s. ForT

0

one reeson or another 911 of the other M?CLVCP 9nd exot ios were

rejected by t1ese authors. There in, loweve , little evidence

to indieeto tbht extensive field t1i9l under forest conditions

in the n01‘1t1’1e23t of all species and Englrfids in the genus were

undertaken by these workers or otlers. Sueh.9 trial would hrve

been tbs logiefl first stej snfi in necord with both tne osigirel

objeeti es of the prijct annd -95 Tererel farcst trre treading

reeownendetions made by the junior antler (Schrgifier, 1037),

Hybridizction work Wes eerried on du3i n3 the 1925, 1926,

and 1027 seasons. About 15,000 hverid seedlings from about

100 different cross com

forent pOQlfTfi were involved es nerents: three white pop 91rs

and five aspens (Section Leuee Duty);s eventeen bleok poylars

and cottonwoods (Section Re eiros Duty); and nine balsam 50p-

j

lars (oeetion Taeqrersoe Sgaoh)‘. Sections Leuooideg Spgch

and Wuran9a Bunge were not involved.

Stout and So lu'einer (1953), with reference to the nomen-

L.Jo

Cleture they used, stete: "Consider9ble study \i9s d-reoted by

the writers to the identity of the various trees involved in

the breeding and to the wstter of the proee9 nomenolaM1J19 for

them but no attewpt will here be made to revise or critically

to evaluate the names already in use or to survey the litera-

ture relating to pOplers."

It would appear that the authors Were probably justified

in avoidir9 9n QtiflC‘fit to "U

revise or to critieelly evaluate the

1In this group were included some "old bbride Stronwl, oelsmn

~ 9 v v

in onaraeter.

(l

J9mes already in use" but it is to be particularly regretted

thst they did not indicate authorities for the scientific names

they did use. Althouh,in most cases it way be possible to es-

sume the pronerauthority and thus the prOper plant, the use of

such synonvms es "P. envuleta" and "P. canescens" mskes accurate.1 . _ é” ~

Initial selection of the hybrid population of 15,000

plants, b9ased primarily on vigor of growth, was made during the

second yesr. Six hundred seefllirgs were thus sele ctJed. The

remainder, and also those needed out by subsequent selections,

basis of 'tteir performance in the iorestetion planting.

Interspecific hybrids were obtained within and between

representatives of the sections Leuce, Aegoiros 9nd Tacemahaca.

Kore hybridizations, however, were wade within and between the

last two sections and sirnifical 3r ll of the 69 plants in the

second or pre-finel selection were from these crosses. Also,

59 of these selected individuals were talzen from the progeny of

17 different combinations-between the sections Aeeeiros and

Tacamshaca.

In most cases 2 or more sister hybrids were represented

among the 69 selected individusls. From the cross

P. laurifolia", a total of 377 hybrid seedlings were

of t.qis num1>er 10 were mnong the 69 slants in the gre—fjnal

selection.

Kany of the hybrids showed remsrksble hybrid vigor.

-10-

*1

Various hybrids, planted as 14 inch cuttings on ordinary farm

land near New York city, attained a height of eight 9Ld one-half

feet during; the first {growing season. bc?xr<~;>iner (193:7) reported

that the "Strath;lass” clone after 7 years of growth in the

13

’x4

‘1‘

”D

U)

H :3

Q;

.J 3

‘21

0field was apnroxinately 7 i “er 9t breast height

and 57 feet tall.

The ability to root from stem cuttings was an important

character selected for among the hybrid HO)Ul”IlO“. Differences

in rooting ability varieC from ttose which could not be rooted

under nursery conditions to those which rooted almost 100 per

cent. It was ouser»rd that many hybrids rooted wore rendil

(A

from stem cuttinys than either of their parents.

Hardiness was also weas an object of selection, and covered

scu h cha 'scierj 813-4.) 93 {Miity to start ;r()\-'~.th early, grow

*3apidly all summer, and to ripen wood and buds early enough to

avoid autumn or winter injury. host of the new hybrids have

proved to be fuli_y Ewa'd;9 in Laine.

Fesistance or immunity to disease, especially the rust

caused by helampsora medusae Thflm. and the various cankers

caused by the renus Velsa was made an important criterion for

selection. Jhe authors reports-d th<m many of the finest plants

with respecct to visor of growth had to be discarded because of

their lack of resistance to disease.

Selection for tree shape and branching to meet the demands

of the pulpwood industry was attempted. It was concluded that

the most desirable form was the pyramidal or columnar type having

a well-develOped main trunk and few laterals. On the basis of

-11-

fl

observations 01

cluded that it was

but could not be s

\

th

n.,.. e pyramidal habit, Stout and Schreiner con-

9 character decidedly hereditary in peplars

elected with accuracy in your? plants.

Schreiner and Stout (1954 ) described their finally se-

lected hybrids on the basis of Vigor of arenth, ability to

root fron stem CthtanS, ardiness, and reelst 1cc to oisen .

-Since the descriptions were be ed on nursery stock no younx

trees, it was impossible to record the sex, form or t11ee, and

character of the trunk. Ten of the clones were assignad hor-

ticultural names as indicated below (Table 1.).

Avermgya “verage

Clonal flame Parentage D. B. H. L'e1_;._jt

Frye 3.

Rumford g.

Strathglass P.

Poxbury ?,

Andover 3.

Geneva 3.

Oxford P.

Rochester P.

Androscoggin P.

Maine 2.0

J!

Duby.

TA.BLE l.

clonal name

Schreiner

of the Sec.

3‘1U:

after 7 seasons

sylvania on

Schreiner

group of

and

several t

9 (inches) (feet)

riwrc 3. laurifolisf .... 6.7 57.0

nisra X E. laurifoliaf .... 5.2 27.0

Livra X P. laurifolia” .... 5.0 84.0

nirra X _P. tridhocar33*... 2.9 90.5

nizra betulifolia X 3.

Lrl(”007“a“ ......... 1.7

b9V]M011caiiX P. berolin-

0....

ensis"..................... 5.6 29.0

naxizoricz-i"X P. t relin-

ensis ..................... 2.9 27.0

In. axir10111ic231.’:X P. niqra

il9nt1e~ensis .Tfl.......... 4.3 3 .5

L~91w091csiL”X P. tricho-

carpa" ........Tfl..... .... 5.1 25.0

’2: l 5"l10-0999X P. bero Linensis

all other of Ae;irosSec. HTacamahaca Spach;

List of finally selected hybrids showing

and nomenclature of parents according to

Stout, 1934 Average diameter and height

growth «t hont Alto Arboretum, Penn-9

well-drained sandy loam (Anonymous, 1940).

Stout pointed out that selection among the

hous and origi_ral hybrids was carried out 1n

I . - ,. . _ To ‘ A . . .

clnxejlzriinéber rHLrS€JHT recrrtueiiwsn. ‘FN€“T Fglflj evrfllnsfp<=d(' o .L - u u - ,

therefore, tnet final jud :cwent of individual PCTlt must be based

inviour in forest plantations. To this end the ten-L finellv

selected Ivorids have been riven wide distribution for field

trial under forest conditions. Diemeter and lei ht growth as

. 2 . 1 _

recorded in a plélfl‘ in Eennsylvania are shown in leols 1.

Of Special intenest, as shown by the above Toole, is the

fact thst P. nifre or it Cf‘

A

i

3

{w J

t‘J.

m r—f-

,les is involved 9s one of the

parents in SiL of tte ten finally selected inforids. In all,

nine different plents are iqvolveo as parents. Three are black

poplars (Sec. leeeiros puby) and six are balsam peplsrs (“e0.

1rTncemm_.eca Snach)o

Pulping tests of tbe Pytrid clones iSOlatec by Stout and

Schreiner have not, as yet, been reported.

Scbreiner (1985), however, indicated that fiber length and

density of wood of the finally selected b7briis have been at

}_J

east preliminarily investig9ted. This author reported that

H.

d 138 been oboe erve i in Pemnlns, as well as in other genera

which have been studied, that fiber length increases With age

up to a masxiznum. The maximum is believed to be rather constant

and well defined for each Species. On the basis of everaee

fiber length of one-yeer-old wood of the hybrids (0.5 - 0.85 NW.)

as compared with fiber length in one-yedr-old vood of Pepulis

treml‘ lWlGS IleCl’lSUX (O . :55 " O 0 57 711117 . ) , tile 1'} L'JDTOidS a‘npea r: to be

definitely superior. As Scbreiner has pointed out, however,

2 additional clones have also been distributed for field trial.

ee Gruschow (1930) for results in Indi na.

53H

Mrb

-15-

Jbrids' fiber length is longer thanT

Uthis is no evilence that the h

their parents ( . trewuloides was not used as a parent) but it

[*0

does indie‘Mt that fiber length of the hybrids is markedly

greater than one of the principle native sources of pulp.

Further DPLllwanPy studies (S(:h - H119 r, 1935) indicated

that wood of the new, faster growing hybrids was somewhat den-

ser than aspen wood grown in western Raine. This is believed

to be due primarily to the prOportionately larger vessel

volume (i.e., rreater per cent of air Space per unit volume)

in wood of the slower :rowing aspen. This concurs with the

observations as reported by Paul (1950) in the case of other

diffuse porous hardwoods grown under extremely adverse condi-

tions. The resultant narrow rings contain abnormally large

percentages of porous tissue, with corresponding decrease in

Specific gravitv and hardness of wood.

Accordinn to Brown and Panshin (1941) there is no very

“10e relationshie in diffiiuse-porous hard'oods between speci-

Jfic nravity, and hence so

V

rength, and the rate of arowth dueQ.)

to (“De (13.1.fo 0)

ed arrangement of pores of proximate size. The

exception, in the case of extremely narrow rings as reported

by Paul, and as cited above, is mentioned by these authors.

Johnson's (1942) conclusions from studies of the re-

lation of growth rate to wood quality in peplar hybridsl ap-

pears significant. This worker found that although fibre

length in individual fast and slow rrowth arrua 1 rings from

frardidenate and P. aoa X P. tremuloidesHi _.__

arents studied.

the same tree showed a marked tendencv for fast growth ring

to produce longer fibres, data from individual fast grown and

slow grown trees did not show significant relation between

fibre length and growth rate.

In general the author concluded that nothing was found

thnt would support a content-on that abnormally rapid growth

is seriously detrimental to wood quality and that breeding

work designed to produce rapid growing forest trees can pro-

pid gronth and good woodceed with some assurance that ra

quality are not incompatible.

Stout and Scrreiner (1934) reported a description of

hybrids between P. balsamifera Virginians Sarg. (= P. deltoides

barshall) of the Sec. Aereiros Duby and P. grandidentata

Michaux of the Sec. Leuce Duby. From this cross, made in

\

1926 176 sister hybrids were obtained. All of these hvbridsIt

were very uniform in respect to the general character of

1e 0.:

.ves, buds, and stems-—tbere being a decided dominance for

nearly all the characters of the pollen parent.

There was, however, much variation in vigor of growth

noted. Vigor of all the rybrids fell below that of either of

the parents as compared with both cuttings and wild seedlings

of the parents.

It is pointed out that ability to root from stem cut-

tings varies greatly in the paren s. The cottonwood roots

fairly well, but the largetooth aspen roots very poorly, us-

ually much less than one per cent, under identical nursery

conditiont. Unfortunately the rooting ability of the hybrids

-15-

was not reported by the authors.

It was concluded that none of the hybrids of this cross

merited use for reforestation purposes.

Valuable hybridization studies in the genus Penulus

1'

I

CD

(‘1'

Cl"

0)

cf

{"3

|.Jo

I3

(lQSfi-a, 1933—“have been reported veriodically by I M,-_

1933-0, l957-a, 1957-b) of the Kaiser fiilhelm Institute for

o A

Genetic Research in e rmanv.0Of a large number of intra-and intersectional crosses

made, progeny of the following crosses were reported as show-

inv exceptional l1eterosis as compared to parental vigor:

(l) P. alta x treiula

(2) P. alts X C‘PGLCTQiS

(3) P. eucaljpntr s X canadel’: sis

(4) P. alba x nigra pv“911JquS

(5) :E. tren«loi'03 X LLtHlla

The reciprocal of cross number (1) above reportedly

yielded progeny of markedly less vigor than when P. alba was

used as the female parent. Wettstein concluded that geneti-

cal differences in the parents were reaponsible. Because of

their dioecious character differert individuals were neces-

sarily employed as parents in the reciprocal cross.

The crosses P. nirra x lasiocarpa and P. resumowskvana— A.— v

x lasiocarpa were successfully made by Hettstein and the

hybrids are believed to be promising for horticultural use.

Intraspecific crosses involving two climatic strains

of E. tremula were reported by Wettstein (1937-a, l957-b).

Hybrid vigor of the progeny fror this cross, as indicated by

the first season‘s height groowth was well maa1ked.

The breeding work ofettstein has not been directed

-15-

U

primarily to the synthesis of rapidlv growing forms but the

development of new forms less exacting in their site require-

ments. POplar culture in EurOpe has so develOped that the

species 3. alba and P. tremulfl have been almost entirely elim-

inated due to their slow growth, crooked stem form, and normally

small dimensions on poor soils in western EurOpe. In utiliz-

ing these spnecies as parer1tsTett in has heped to obtain

hybrids showing heterosis and improved form but still making

the low site demands characteristic of the parents.

heimburger (1950, 1940) of the Canadian Forest Service

has carried on extensive bre din. Sthdj in the genus POpulus.

The ObjeCPt Of the work has been to produce hardy and disease

resistant material of vigorous growth for the produciion of

wood of high quality to meet the demands of the match, veneer

and pulp industries of CansMd . In addition some attention

has been dir cted to the develo;1ment of material suitable

for sLelterbelt plaantings in the 12rairie provinces.

In 193d uci1burer reported the successful crossing1 of

$.21

VL‘

P. alba L. L P. carescens Sm. (probably P. canescens (Ait.)

Smn)2 with P. tremuloides Lichx. and P. grandidentata richx.

The cross P. tremuloides x irandidentata was also successful.

All of the above parents are included in the Section Leuce Duby.

Good seed from the crosses P. aloe x tremuloides and

P. canescens X trewuloides were obtained and plaitcd but soona-..

after germination the seedlings perished thru dampirs off.j\J

1"'etts tein's "grenhouse teecunie_u_e" ofusing cut twigs as de-

scribed p. 67 has been used exclusively by Heimburger.

Rehder, 1940. This plant is considered by Neimburger, wett-

stein and othersto be of hybrid origin (P. Elba X trefiula)

but is not so considered by Rehder. —

Seeds from other crOSS:S were planted in ric1 sterilized

(formaldehyde) garden soil covered with a thin layer of med-

ium fine 1.’1'*1sl‘1ed sand without iurt‘rler loss.

23

5

l

PI‘O 5VJ“;T of E, “JAN1:X 5rsndiderimfiyi, and E} tnnymiloides

x grandidentatal uniformly displayed characters 1n|o°mcdiqte

between the two parents involved. Pro5eny of 3. canescens x

grandidentata, however, showed great variation in vegetative

characters. It was concluded by leiW¥1r5er that this would

be a normally antic

character of P. cane

ioated result due to the presumed hybrid

scens. This appears to be contributory

c oncl lsive eJOvut not

BGWMHUWzBP lat

ity of root-cuttin5 s

brids.

made in the spring

hybrids and planted

at this time because

aspens indicated that

insure

tte unique idea of t

by this method.

author to be a 500d

climate.

Interesting

All theof 1957. g.

XL

At the end of the first

of

establishment.

Survival of the cuttin5s

survival results

.randidentata clones

vidence of hybrid origin for P. canescens.

er (1940) reported tests abil-

Of h“\f a-

U

1- ' J- A L ‘

a ”8F51h the aboven6 3

5rowin5 season (crosses were

1956 were made of the)

in the nursery.

root cuttin5s30

The cuttings were made

previous investi5stion with the native

the cuttings must be fully dormant to

Of (rests: “nificanoce, however, was

estin over-winter hardiness of the hybrids

is believed by the

test of its 5eneralaslaptation to the

shown in the summerwere

tremuloides X grandidentata and P. alba

survived but a number of the P. canescens

1Victorin (1930, 935) reported rstuial kybrids of these

crosses in Quebec. qUnder normal conditions in Canada, how-

trWIlo.d flowers about 10 days before P. srandi-

but at aDCiT the same time as P. alba (E1eimourjer,

Natural hybrids of tre let t6? havebeen onunorc

with the Ortificial and found to be identical (Heineu; 5er

1940). Tubeuf (1(351) reportsd natural hybrids between P.

alba and P. tre~ula in Canada.

X {unnsdidrn1vvts ne:drfi<yi. Suclllml“"lOLC)l'“1l vaTq_Ition W98

antic pated on tne ‘asis of the wrest morpiolo5ical vari_stiq1

‘5

(I:

H O C U)

H

’3

‘i

'J.

{3

Q4

H.

O «N1

rr

(3

0..

w 3

'5 JJ

"D

Followinr this natural Selection, resistance to poor

07‘) {Mid cinx3-1)a(fi: (BQLESGKl tvw Twiifi leldjllfll SI).

were artificially selected for durinn tke lHSe-R? seasons.

x *rnr’midet9 1‘8 1:3'LPE In. T‘LGSE) l'v’. r‘ifis '3' Owew vrec'; ; 10 "~00?"

5roztr and great "‘sceptibilitv to die-buck tut, on the other

Lap“ gio'ed a qur dsugct L: lasistrn e to u819TPSOPa op.

rust. Heimburser su55ested that such extreme SWHSCEDUibilitv

‘

of some hyhrgcs to disease may be added to the mecrenisms

of isolation which reintain syncies differentiniior as de—

Ir‘ I" - '0 N. '3 "“"

Snitcsecgierit Iiexue

clones 5ave pronise of valuable varieties available directly

. r- 2 ° v -.". » ---" ' W» - ".~ x- ° , w

for reioreststion 3l_w21158. ”ell-marked :Jtrid V1003, easy

prepsgstion by root cuttings and Hell~ defn‘o. resistance to

Helempsora rust and r-;_

About two—thi: OJ

L]

O W C‘?‘

:3”

W}

O p-

LJO ’

'Jo

:3

f.)

l...’

Po0 O T)

U)

’3

C‘

d)

N

1

1‘

J 4 .J

(3

LJ.

I

dentata progeny were eiscarded by :t; end of the 1939 season

but the selected plants are sail to represent some of the

?8Rd. Some were superior in the

various selected claract rs to the P. alba x 5randident9ta

hybrids. It is planned to reneat the ori5inal P. canescens

X :ranlioentota cross on a lar5er scale, using the best P.

Additional hybridization nor? ctrried on nnring the 1957 and

1(38 seasons .ws reoorted Ly1e1m01ror (1940). bince 2;

91‘ o I, an? its ‘r13‘b13‘63 ,1 176-1 I’Qiufltedlj,’ 1OO'C‘361 a

marked de5ree of drought-resistance (Hettstein, 1939) being

used in shelterbelts in southeastJrn

*ubbard, 1956),

trewuloidesc_ba x P.

ori5in was from native stOci

prairies, taxonomieslly undifferentiated,

(Moss, 1932)

and.i_n erodimywi‘rOPlslss. AJtTTV115h_ ttua cro

it was ncluded that this was not die to

poor technique.

The following crosses were,

P. canescens x P. trrwulo1dps (”egte in")

alba_\ntr. Bollesrrtlxn1che (0‘

(UV, ()8Ste-1,171"

however,

the

P. trewuloides). 1“n---‘-J

northern

b1 1t reunited 11* more

other native

Wes uniucce sful

iroowr‘tibility but,

reported MUGcessful:

Rehder, 1940); P.

nranoi03ntata)l X P.

O

‘13

1...:

OJ

’33

>4

aniels f. (rt—

A. i 7‘ . \‘~ J- (1 1‘ (1 T‘ v

5rsn(ident ca) X 5, 011080cens

x grandidentata)l X E, nutenei (or: P.

sures;

(triploid)2; (g. alba

canadensis Eugenei

(§imon-Louis) Schelle, the "Caroliz1a Popla

P. trenuloides x P. adenOpoda Paxim.; P.

. . 2 . w

(trip101d) ; P. ocuminata Rydo. X P.

Rehder, 1940);

tremuloides X P.

canedensis

alba

T

r3u5enei;

X P. berolinersis Dipp. (i.e., 3. laurifois x P. nisra var.

italics, Rebder, 1940) x x "NorthWas t Poplar" (sirilar to P.

Jackii Sarg. whic}1 is a h

a. deltoides, Hgimluu'iger, 1940).

ybrid of the cros

iA natural hybrid, exem1ned oft faogically by Pete, 1958.

“A triploid form, examined crtoloinCally by Peto, 1955.

-oo-

Of special genetic interest in the above cres, s is the

nrelininary ana1331s of the mode of inheritanoes of branching

habit and leaf form indicated by the crosses: P. canescens X

P. alba Bolleana Lauche and X P. bero1irensis Dipp. x X "Iort11-

west P0p1ar"

Only two YNotional nroesnr of the former cross were

early selected and prepa5atsd. Jhese two plants late", however,

proved to be of considerable interes . Loth_slowed the ”ex-

. O 1 ‘. _ 1- - -‘. V -

cess1ve Juveaiile drenchineSs' craracter but ne of the seed-

habit of P. alba Bolieana,

CD

d (D

3 3 I) s; to

O (73

H U)

0 1.4.,

Wcombined with the leaf and o C3HQSC€FS¢

bThe other seeclina had the normal, divergent branching stem

character of P. alba L. combined with leaves more closely

res embling those of P. Cine cens. Althou15h scanty evidence,

it does at least indicate that inheritanc of certain branch-

1

ins and leaf characters (lobin5) does not involve linkage.

Since both parents in the ,ross X P. berolinensis Dipp.

X X "Iorthwest POplar" are themselves hybrids of a Section

Tacamaraca x Section Ae5~circa cr as, their progenv siowed a

wide variation in Vigor and leaf and stem characters. Of the

26 plants selected after discarding some duarf seedlin5s, 12

showed the "exces ive juvenile branchiness" character, charac-

teristic of X 2° berolinensis, while the remainin5 14 had the

H

"clean stem” character. Heimburger concluded that excessive

juvenile branchiness" of X P. berolinensis was dominant to

"clear stem" :no way be transmitted to about one-half of the

progeny in this cross.

1The cross probably represented a back-er ss as far as this

reticular factor is concerned.

-01-

O 0‘ _ ._’O K ' -"..\P (I. O '3 F'.‘I‘; y t"

Excessive Jivenile branc;1ne s is an uLueclfaJlC charac-(,0

tor for the production of clear wood for various industrial

purposes but for shelterbelt material it is exceedingly de-

sirable. For this reason the shove cross has been repeated

on a larger scale by this worker and other "Korthwest Peplar"

material, more resistant to Septoria canker and Melanpsora

rust has been employed. Results of this work have not been

published.

The two crosses involving triploids (as listed on p. 30)

were reported as producing few seeds but the hybrids have not

been studied cytolosically. The cross involving the canescens

triploid produced progen¢ of promisinr appearance which have

shown reasonable vigor and freedom from disease. All of the

crosses noted on page 20 were aralvzed in considerable detail

by Feimburger, with particular attention to disease resistance,

and form a valuable contribution to the genetics of tiis im-

portant enus.(1'‘_

V

Bogdanov (1954) concluded that it is only possible to

graft pOplars success.ully according to the series: aspens--

U) ilver pOplars—-cottorwoods-—halsam peplars. Reimburger (lQAO)

ointed out that this sequence agrees With the series of suc-

’53

cessful crosses made by Yettstein (1933-2) and Stout & Schrciner

(1955) and concluded that this strongly indicates a similar

series in genetic affinity of the different poplars.

Since BOgdanov (1954) reported a successful graft of 3,

nivra L. on Salix viminalis L., Heinburger (1940) attempted

the cross Salix virinnlis x P. canadcnsis EuQenei (Simon-Louis)

Schelle. The resultS, however, were negative.

(U- ‘1‘,-

Swith and Nichols (1941) have repor,ed what they tenta—

tively believe to be successful crosses in this genus. Of the

26 cross combinations reported, the authors indica'ed that the(—h

4 crosses involving P. raX1Hoticii Henry seemed most prowis-

ing. Comparison of vigor With the parents or expression of

other characters in the hybrids have not as yet been tested

or reported by these workers.

”ilssor-Ehle (lQBE-a, lgde-b) and Bergstrom (lE40) re-

ported genetical and cytoloaical studies of successful diploid

x triploidl crosses of E. tremula L. The crosses were made

prrimarily to determ 1ne thepossibility of obtaining a tetra-

ploid since hfintzing (1956) reported some giant pollen grains

(with possibly the complete 3n complement of 57 chromosomes)

produced by the triploid from Shane. Interest in securing a

tetraploid was prompted by the presmted feasibility of produc-

ing triploid seed at will by diploid x tetraploid crosses.

Of the approximately 100 plants obtained from one of

the crosses2 (involving the Shane tTJImlid clone as male parent)

the majority were aneuploid (mostly hypotriploid), several

were exactly diploid, l exactly triploid, and l exactly tetra-

ploid. The other cross involving the Medelpad triploid (i.e.,

from the province of Ledelpad, Sweden) as male parent pro-

duced about 20 plants,practically all of which Were of poor

develOpment.

All of the prooeny of the first cross proved to be

healthy vigorous plants. The triploid and near-triploids,

however, were distinguishable frori the others by their ex-

1 Triploids were of 2 sources: 1 from province of 51Wape, south

Sweden; first reported by Nilsson-Lhle (lgbo); the other from

MedelPad, northern Sweden («icmnnl t, 1937 . ,Tettstein's greenhouse tec'qicre we3 ’used fiettstein, 1929,

10-212 1«\ “'2

tremely rapid hei ht growth. The tet*aploid fell somewhat

below the triploid in vigor but had tie largest leaves of

all the pregeny. Tue intermediate neuploids showed poor

development and were 1ihly leCPWSAU in appearance. iMaILy

died at an early age.

,0;

Further work with the trioloids an tetraploids is re-

.

ported in pregress at the Institute for Breeding Forest Trees

at Svaldf, Sweden but no further reports have as yet been pub-

lished.

Jornsson (1940) reported 01038 3 of diploid x triploid

and triploid X diploid E. trewula L.1 and their cyt0105rical

analysis. A totil of 29 crosses fielded about 600 plants...

\

0

The cytoloeical results were for the most In:rt similar to0

those recorded by Eergstrom as noted above, but triploids and

tetraploids in the pro rny fell far below the diploids and

even some of the aneuploids in hei ht romvt The author,

however, concluded that the cause of these differences may

be traceable to the occurrence of photoperiodic races within

3. tremula vmlich may be critically adapted to the special

light climate of their habitats. Since the tetraploids were

chiefly offspring of one parent not native in the area where

they were grown (Svalbf, Sane, South Sweden) it is believed

that their absolute vigor could not be accurately judged.

Several workers have reported miscellaneous hybridiza-

tion experi1ents with various members of the genus. Gambocz

(1926) in Hungary and Albensky & Delitsina (1954) in Russia

reported successful crosses between P. alba L. and F. tremula L.

l hale triploid from Shane & pedelpad (used also bv Barbotpom

1940); female triploid from-. orbotten.

-24-

n _,,QThe latter workers, using the reeLhouse technique”, obtained

of 62 plants which demonstrated a

very marked variation in morphological characters among the

individuals. Visor of growth also showed a nuraer of grada-

tions. The pOpulation could be rather definiely divided inn)

a so-called dwarf group and another group of very luxuriant

growers.

A

Re"nier (1934) reported an csscntiall* taxonomic study

.1

of a large number of peplars collected in France for breeding

research in diseas» resistance. A few preliuinary notes on

the reaction of various species and hybrids were cited by the

author.

Vloten (1958) made a first report on investigations into

1

the susceptibility of peplars to Jothicniza pepulea Sacc. et

Briard in Holland and other countries. From.comparative ex-

periments made at various centers in Holland, but especially

at Hoog-Keppel, it was observed that peplars of the section

Tacamahaca (Balsam peplars) and the hybrids of Stout and

Scb'einer were more susceptible than peplars of the Aegeiros

(black pOplars, cottonwoods) section. On the basis of these

obserVations the author sugvested that Ae*eiros ionlars shouldV \d A ‘.

receive preference as parents.

Vlotcn noted that P. marilandica Bose.1 and P. recenerata

Henry (or: X P. canadensis regenerata (Schneid.) Rerd.) ex-

hibited a high degree of resistance to both Lothichiza and

\T

i: e O ,D

tri- canker. P. brabantica Houtzagersl also dewonstrated

high resistance to Dothichiza bu d-

.Jo

was found susceptible to

lxot listed by Rehder, 1940.

.3

\J

Nectria and "bacterial canker". The author concluded, however,

‘

that no peplar is completely immune t Dothicniza populea.

Hvbridization in the "enus Populpfi is consiedrably fa,-

1itated Ly the usual dioecious condition of the individuals.

:eater impeltance, however, is the fact that success-

ful breeding work may be carried on in the greenhouse usins

cut tWiSS accordinr to the method orioginally described byr

U

Yancnevs‘i (1904) and later perfected by Wettstein (1929,

1953-b) for Salix and Pooulus. The method has been reported

PU...“

successfully used by Al'bensfiy & Delits na (1954), “eimiurer

(1936, 1940), Berstrom (19‘0), Johnsson (1940) and Smith and

Nichols (1941). Jablofikov (1940), a .uss ian vor|er claimed

that the type of hybrid pro:luced by crossing trees growing

in the open differed from those produced from crosses be rformed‘—

on cut branches in the laboratory. ]t is probable, powevr~’

.1

that the variation observed oy this worker is of a simple

genetic nature since the Species of this genus seem to be

characterized by noticeable hetesrozy0:;i'3. The wide varia-

describedfltion between individuals in groups of hybrid fro ena

A. \J ((4

by Schreiner, Heimburger and othe33'would favour this conclusion.

Heimburger (1936) reported the successful use of cut

twigs sent from considerable distances and sugyested that

this nethod Opens up almost limitleSu pos mililities for the

q

tree breccer regurdless of

7

is location.

Heimburger (1940) in OOWTT ent in; on the “reenhouse method

of hvtridiz Miin reported difficulty in getting seeds to nature

-26-

on {watle P. “"endirientsittlric heiu.. tun: s. Thyttstedgl (1997$J))" '_ I- :- .84_

also resorted marked differences be,wern vcrjous specigs in

~ 0 0 0 _ -‘ ° ‘», - .. _‘nr’

their atiljty to set setu in tun 'e~ ousw 0*( lttc: (1931-b)

- .. - - 4 v - :° -,describrd inFIV1drfl ffe">zces tithln tie 31~c species Xiii

,_ o _a v. 'I ‘V f’\ a, ~ ~ ' .1.. J_'I‘ J...” «‘7 ‘ _ ‘

gri fl,& Licnois (lJil) had success city tge «reenLOLSQ\a'

w_ '1 o Q _ . > O - . . .> h 0 J_ ‘1. _‘ V. r J_ '

refinOQ only in tne see 10H Leuce. Crosses in ObJUP seenions

of the genus were more successtl when nede on the tress.

Forcing of pollen however w~s used to sdvaute;e by these

workers With $180198 from all sections With wricn they worked.

(Vi-1‘ " "\.""-" ‘ 1 ‘u (71-:- ‘\

on}: in: kb'lb ICA; 1U

" O "‘ ’3 ‘1 .. F» 2'. r '3‘. ‘ ‘. .. C . w

as iL-s (1910), nrisnson . Le mans (1997),:loensrr

(1940), Peso (1956), and, doubtless, others have reported

members of the genus showing a monoecious or polygemo-monoecjous

condition. Hestings Observed a peeimen of repulus grandi-

denteta Micheux for two successive veers Which showed abnormal

flower develOpment. The tree was essrntisllv pistfllste but

several branches bore cetiins whicn Were nude up of male, f9—

msle and per set flowers.

The abnorrnel I‘. tremuloides Licheux reorted by Erlerson

& Hermann had about 82 percent perfect flowers. Ore percent

of the flowers were e3: lusivel.;r stsmirete and 17 percent were

pistfllete.

' 0 1 0 a w 1 o

Albensky (leO) reported sn instance of ue‘mprrooite

flowers in the aspen which was not inherited by its In0Qeny.L O

In another instance, nine hybrids of a cross: aspen X wLitmJ

I" 7 J.

_Lpoplar were Ciro 'xllly stu(i ed as to their sex 8X res:ion. One

of the hybrids bore male flowers only,four bore only pistil-

te flowers, two failed to flower, and two had flowers of

02)

both sexes. In the latter cases, whole catkins were male or

female or scattered catkins had both male and female flowers.

erfect flowers were not observed. :nese monoecious trees

’1!

showed normal develOpment of pollen and fruits with sound seeds.

It is probable that sex modification in the genus occurs

more frequently than the literature would seem to indicate.

The transitory flowering period and inconspicuousness of in-

oividual flowers may mask more frequently than is int

a so-called abnormal sexual condition in this genus.

As pointed out in the case of Salix and other dioecious

plants, the occasional occurence of monoecious or perfect

0\

flowers may be of assistance in providing a UOSsib lity forJ.

true selfinv. Accordinr to East (1940 the occasional )er-0 u 1

feet flowers in this genus annear to be self-fertile. Such

a condition is obviously favorable to the breeder.

POLLEN LOEGEVITY

Studies of pollen longevity in POpulus, reported by

BOQGanov (1955), indicate that refrigeration at a tempera-

ture between +5 and -50 C. may be successfully used to store

pollen. Pollen germination tests of fresh pollen conducted

by this worker indicated that germination percentages varied

greatly from spec;es to species. For this reason it was dif-

ficult to predict exact lenghts of time for successful storage.

P. laurifolia pollen, having a high (QOfi) fresh pollen germi-

nating percentage, was successfully preserved for 15 days under

-2s-

the conditions described above. Pollen of P. sucveolens,

havin. a low (35;) fresh pallen germinating percentage, ger-(T

u

minated up to 19 percent even after 45 davs. Although tercent”I

of relative humidity is not mentioned in the abstractl of thisf—f—

article, i1 is assumed that it should probably be waintained

at or around 505 for successful results.

J- 1

It would appear that the pollen storage eecnnique used

by Reimburger (1936) should have certain distinct advantages

over the customary techniques which must be used in most of

the other forest tree ggnera. His method consists simply of

cut twigs bearing wale flower buds in a snow-bank,

or refrigerator, and forcing the pollen as needed. There are

probably time limitations to this method which might be ad-

V‘GETATIVE PEPEULUCTTON

There appears to be abundant evidence of a more or less

empirical nature that most members of the genus Ponulus may

be reproduced readily from root cuttings. Baker's (1918)

-n and‘._.lo

studies of P. tremuloides iichauxg in the Gpe TD

t Bas

the observations of Kittredge and Gevorhiantz (1929) in the

extensive aspen areas of the Lake States supply evidence

that both P. tremuloides and P. grandidentata apparently rely

more on their vegetative reproductive ability by root suckers

than on seed. The frequent fires that swept over the lovced

over areas in the Lake States are believed to have been the

chief contributinf factor to the denseness of the present as-\J

iSee literature cited.

“Probably the Focky Lountain form (P. tremuloides aurea

(Tidestr.) Daniels).

on.v 'a

pen stands. Althouh aspen is readily killed by fire, the

protected roots send up numerous suckers folloving each fire.

By this means sinulc trees Lava esta7lisEed large clones.

The rootin " abilitv of stem cuttings apparently shows

the various 5.

fr)

(3

K)

H.

C.)

C)

E .5 ‘4 C

4

.1

H 94

U)

H

considerable variation amen

varieties, and forms in the genus. host workers (Stout &

Schreiner, 1933; pe1MUur’eL, 1958, 1:.40; Sno , 1958; and others)

seem to concur in the belief that as a rule tne assens and

white poplars (Section Leuce Duby) may be grouped as ”dif-

ficult" or resistant" plants, wremns the black pOplars and

cottonwoods (Section Aeaeiros buby) are considered easy to

root from stem cuttings.

‘Eith respect to one of our native spens at least there

are, however, conflicting points of viex. Snow (1958) re-

H

ported that nder norm7l conditions dormant cuttirn;s of

either large—tooth aspen (P. ”rancident‘ta ulCJQUX) or

trembling aspen (P. tremuloides michaux) are difficult to

prepegate vegettatively,se1dom rooting more then about 1

percent under nursery conditions." Thimann & Delisle (1959),

however, state: "As is well known, cuttings of aspen (POwulus

tremuloides) and Lombardy poplar (P. niura italica) rooth

adily .

Such diametrically Opposed points of View with regard

to rooting ability of P. tremuloides discourages any specific

or even general conclusions with regard to the rooting charac-

ter among species or sections of the genus.

Snow (1958) was primarily concerned with determining

tie best tine during the dormant poliod to make cuttings, as

well as tie best concen13ration of auxin and Optimum period

of treatment to use. he concltded that (l) dornant cuttin5s

0f bOth P. tTJFUlOiflGS and P. 5rendideateta can be rooted to the

extent of at least 65 percent by treatins with a 10 m.5. perK.)

,ietlflh of 11roleb1tric acid for about 27 hours;

and (2) maximum rootin is secured when cerwawt c1”t7“s are0

Q rn

\u

U ‘Z$

L10

:3 l

‘made while the buds are swellin5 in th.

The work of Ihimenn & Delisle (1939) with P. tremuloides

was not .esigned to dcnonstrate that routing abilitv in this

species could be improved since they assumed it to be well

known that the species does root well. Significant, in sup-

Jort of their contention,was the fact that untreated controls

0'

ll produced roots within a 5 week period. Cuttir —— ("fl (7' ‘7 .3 ‘4

ID, h.O"’e'LI ,:13

With varyin5 concentrations of indole-E-acetic acidcf-

C1—

0)

Q1

res

were found to have an Optiw111l concentrai:ior of 100 he. per

I

liter jud5ed on the basis of number of roots .roduced. leis

.1

Optimum, however, would not i7rove Optimal in practice

pointed out by the investigators, since e?Htc1e inhibition of

buds was demonstrated by t}7e cuttizigs thus treated.

It is worth notin5 that Stout & Schreiner (1953) reported

thtt rooting ability of some hybrids showed great improvement

over rooting ability of eith1er parents. Specific cases were

1

not, however, cited by these workers. It is conceivable that

such obser'at:1ons may be greatly influenced by the a;e of

stock from which cuttings are made, the inportance of Which is

emphasized by Thimann & Delisle (1939).

lvical characters of the Various pOplars and cul-F10

The s

tural practices used in forest plantations in several coun—

tries have been descriLw-Jc “23v mrious authors.

0 - .‘ “ 0 ‘ - l r ‘. 0 _- . . -

Forestry Connissior Pulletin to. b (1923) 18 reported

by Cansdale (1939, to provide a usefil sum sry of cultivation

methods in Great Pritain. French practice is surp nrized by

Breton-Bonnard (1902, 1929) and interest in rcplar cultivation

in Germany and Switzerland is reeflected in the publications

\

by Leiber (1936), Zircber (1936), Bar1e3 (1936), and Hettste LJ.

*1

(19W7d, 1959). heeiLm (12)SO, 1931) has summarized Dutch

practice. fieigel and Frotringhcm (1911), Buttrich (1921),

Kittredge and GeVOIEiantz (1929) and others have described

CUDOKOSCFE RUNNER

From lists of *‘onosole nuxoers and other reports nub-

1ished by various authors (Blackburn & Harrison, 1924; Meurman,

1925; Gaiser, 1926, 1950-b; Tischler, 1927; Er arson and Her-

man, 1927; Blacl-zburn, 1929; Ifintzing, 192363; Ilakajima, 195 ;

Peto, 1959; and Kaude, 1939) it would apxesr thvt the basic

number in the genus Ponulus = 193 .

Lillewi jn (159-b,1940) believes that Populus (and pos-

sibly Salix) is a secondrry noluL101d vith an original basic

number of U. Four of the orismi91 8, according to his hypothesis,

have been triplicated, the other 4 duplicated, and the present

lGraf (1921) in an article consisting chieflv of a morphologi-

cal description of P. tremula L. and other specie s, first

reported chromosome_nunoers in the genus. Hias count of n = 4

for P. tremula L. does not agree with that of later workers

in this or other Species in the genus.

-52-

.J J

4

J -_J

\

r-‘ d

'_.J

L

_T, was t%e resil’ of s fisiow of tuO CVPowoso es Witn-r

-. "I_1 ‘ - ;_

one of the greats of ttree. his COHCLHSiOIS see uflSCd on secon-

GHPYd'""OCin+io1 of chromos mas Lt wetnnhwse I.L -

YVWIDTHTlV

31-13111; -UJ. --J\’.LJ-

‘lJ*ovx)Lrll(F ER}? 6. orx (1 M44) rtgyartnzd 21. l :lssuiifvgra QL.

(= E, CN3lf01588 fidxxmfzrieysis imxxry, V9?. ) emsza proheiflxg detr9_

PlOld (?0 i ') Dot “JHZ(1TSS ripoyted it as a normal

1 f‘0 O ‘ ' - _ I. I ‘. - r': "‘1 v7 0 4” 1r .5 ‘ x T I V“

d19101d (2n - so). As p01nteo out of allo. gnu “10M 0 H U)

A

H .g

H

v V

' M , 3 - ~_>_ ‘ ’ -___ 'a o :1

howevee, it Ki? elso CO”CGquUl€ OQRQF in tne tetpwnlOir form

-7 _, _._ . .o .. . -. .A ‘. _ p. I. 1

in nsture. In tge swno yVyeJ, lenruan recovlod toe reouoed

a _ ‘ -. . 7 s '7‘ N ._¢-" 0 o _ _ o 7‘ . 7(‘ 1‘ 1 - -

CILWYH0¢0W€ YUP”??? 01 17. bt:ofififi.fkm“rlere ass.)d eno t;xn;¢1 tetra-

--oid. Later counts of CEI‘I'IOI-I'Iomarne mmbe‘r‘ in tie-"7. species by

other worgers PQVe not cove to the author's attention.

Naturally oeourin; triploio or Approxiwntely triploid forms

Ol ix”“” "Peeiws in Lhe swvtion Lenoe Duuv lene r1 Femorteo

in recent years. Lilsson—Ehle (1930), Ploomqvist (l W), and

lelsnder (1933) all reported finding strains of e ”Qigws” trip-

loid (2n = + 57) form of P. trwnnls L. in the forestSof Swefen.

, Kilsson-Ehle (1935—2, lofib-b) and Fobnsson (1910)

'1 _ ’ . ‘ -'- - ~ A‘ -. . I' . .1 . 1 r‘ .I- -\ :1 . -‘ . f . '

rammieteo S’THAJQSlS (Hi tEDldeLOLQF, 'tlelCHflo wiml Vflfjxflxs JILLQP-

mediate aneuploids from artificial triploid X diploid crosses

of P. treale.

Peto (1958) has reported OPLH“l€PP' of triploid forms of

30 811351 1.1. 81’1d Po 091-9'71918 (fiiton) SII‘W introdilccxi in CQqng‘.

DillgI.jijn (1939-2, lUBD-b) also eeported a heturally occurrin"

Tr oid form of P. slbn in the Netherlinds. ”nthsbein (1973-0)

reymnfixfli'Peorooertqtnifios of P. elta.ln iirnsstithed i11ik3wnany—- \J

88 normal diploidy’ It 13 probable thfit bOth dlfilfili SUC tri

ploid forms of this Species exist in nature as has been con-

cliusivelfir(Manonsiauatci ill the (rise ol'E’. trerafiti L.

CYTULITUIXUL ETULIEL tfl’fEKhClht, lflfiiflETIES, AIHJEIVWPIDS

P. trewula L. diploid (2n 3 5b)

Blackburn and Harrison (lf24), in England, studied meiosisl

in pollen mother cells of the species (also 3. niera L.) and“I’—

first reported the haploid number of 19 and diploid nunber of

58 in the genus.2 fietustein (195m»- ) in Germany, and Rfintzing

(1936) and Johnsson (1940) in Sweden confirmed the chromosome

count for this species.

heiosis was reported without eppe rent abnormality by

Sleckburn and Harrison. Chromosome morphology, described and

illustrated ov these authors, is said to possess characteris—

tics sufficiently well-warked to enable individual identifica-

Qtion of the chromosomes. The diploi chromosome complement shaved

10 pairs of autosomes and what was belileved to be an unsoual

pair (heterocrromosomes) at metaphase I. The pairs of autosomes

ranged in size from one large or giant pa 1r thru two varying

sets of medium-sized small paris. With reference to the heter-

ochromosomes, the authors concluded that "some evidence exists

‘

oossioly sex-determining5.

A.

of the presence of heterochromosomes,

c, in the male of P. trenula."

5fintz'rw (1950) found meiosis for the most part regular

lFixation: Carnov's fluid; stain: Neidenhain‘s iron-alum-haema-

otoxylin, W1tUOdfi COUHteP Stfiin,

“Studies of Salix species as we 11 led them to the conclusion

7tth 19 WES—BEETC number for the fonil Saliceceae.

oChrome-acetic-formalin

fixation with short preparatopy treet-

meflt in C 1! P1103.

P;

in normal 3. trenula L. diploids and concurred with_the morpho-

ical description of ctromosomes riven by Elackburn and Har-

rison. This worker did, hovever, observe that occasionallv

txo univalents were observed at metapiase I in some specimeno

studied. In such cases there Were observed 18 bivalents + 2

univalents.

Johnsson's (1940) study of moi sis1 in si;teen 3. trewula

L diploid clones fror various provinces in Sweden sh Wed 12

which behaved quite norrelly. The other 4, however, showed

from 19LJo

iilarit*u (3dro»oscwwa'bodies rern it“J x.) &

bivalents to as many as 7 bivalents + 24 univslents at netaplase

I were observed. In the latter cases , the univalents were dis-

ributed irregularly in the cell and divided or did not divide

at the first division. The svlit or unsplit uniial nts were

either included in, or left outside of, the interphase nuclei.

At division II split univalents included in nuclei passed toA

one of the two nuclei without further diviSion. mhe split chro-

Insomes excluded free the interphase nuclei were able to finish

the second division separately.

Johnsson also studied pollen duality and found that theJ. l

12 diploids showing quite regular meiosis produced a mean aver-

Dage of 79.9 per cent food pollen. Percentage of good pollen

among the 4 clones with exceptionally irregular meiosis varied

betveen 38.9% and 80.5,A. The author concluded that variation in

pollen quality in the diploid form of this species can to some

extent be attributed to meiotic disturbances.

h short breparatory treat-A

10hrome—ace tic- fOIIn::lin fixation wit

ment in Carney.

"Titll'rezvird. to £3 c ()cczukrernge (3f LU"i\fiMleIHZS ill di.‘

1.

Johnsson sv“‘csied tlwt temoereture chsnves in early' .L ._ ' U

leztotene end oachytene In Gistgria trigona :f‘30tcd *hc chiesre

ne ”'Plly with ligl frwowoecy in *snv cjoloiis of His syosi,s

an”, in t‘erz, furnish ()LQL-L‘wt-mitlet: for U_ured‘-.‘»_ced (2n) gfl'rzietes.

This conclusion is suggortcd h; the foot ttst triploid 5. tre—

rile L. is lOt especially rave in nature.

2. tre ulq L. trijloid

'Irc flhfllnu of the fins; trijloid cloze of E, tremula L.,

rdcr LillB, Sigre, Qout: Liedtn, ‘s rejirttd ‘3 Yilsson~tule

the etude tine (1"‘331tzih tc- JOhIlSson

C)

”J

0 1’3

CL

Ho

:5 1

19301. By 1940, d.

(1940), eight wore clones had been found: 5 in middle Sweden,

5 in north S eden (Ploucvist, 1937; Tonetorp, 1937; islander,

'1 _ v

U930 triIn no cese hes tnc somatic number for C‘t

loids

Fa)

been ascertained to be exectly 57 chromosowes, but rather + 57.

)

nd especially

\1

Nilsson-Ehle (lSSd), Lelander (193$ , and Tometorp (193

-10

(Dfin

-Loreported thrt the apparent vigor of tlc triple

the larger and darker leaves were resr isihle For attention

being drawn to them. These chindcters were co seouentlv use.

as criteria in huntine them.

see also Spregue (1940).

Lititzixt; (heteivnirwni Tflue EUJeIWJCfialeThSE?! oi' stcwvata.:for"the

triploid from Shine as 10.76 + 0.13 units and 6.5d + 0.14 units

—‘

for the normal diploids in the vicinitg. Leiender reported the

averunje sixwuit; lullfth as lele + (?{13 urdflna for CHM? of tiwa tri-

oid clones from horrhotten and an avers e of 6.36 + 0.15 for

in oi ltifll stovete lxuwgths CL1 snail-lfirivvi(ti,loi€3 :ru, lar e-- \J \ L

leeved diploids and triploids and concluded tth, usfng sto ate

1 O ‘ _ I 3 1

lengtn as a messure of cell Size, 'the Size oi the leaves oe-

sends on the size of the cells. The size of the cells in turn

deoends on chromosome number (triiloids having lsrger cells than‘ A

most fiiploids) and on other prisuhnhly hereditary factors not

known (there are some diploifs hQVing es large cells as *he tri-

ploids and even larger)." Leaf side is, acoordiig to the author,

nevertheless of value in searctiwg fer triploid aspen since it

appears unlikely that small-lesved tr'ploids exist.

Ldntzitg (1956} and Tometorp (1937) reported thvt tri-

ploid forms of P. tTJHUJQ L. have larger pollen grains and poorer

.Jo

pollen than diploids. Lflntzing obtained a definite bimodel ris-

trihution of pollen but did not give absolute values, Tometopp

on the other hand, recorded poll 0n diameter from 2. microns to

42 microns. This is comparable to triploid pollen diameter

ranges recorded by Peto (lESo) fo: P. lbs L. which was not con-‘QD

sidered by Peto to be definitely bimodal.

Although egreein th t, on the average, triploids have

larcer pollen grains then the diploid, JOhnsson (1940) observedU U

no "bimodal curve for the pollen diateter of trinloids" as re-

ported by Fdntsing and concluded size alone could not be used as

-57...

a criterion of triploidy since tbere is a significa.nt overlap—

of size. Johnsson, however, concluded that pollen sco‘le U)

have a limited value in distinguishing between diploids and tri-

ploids if both diameter and quality are taken into account.

The author observed ttat triploid pollen, in addition, sometimes

shows "one or other elongated, sometimes constricted grain".

Peto (1958) suggested that environmental influences may

have been responsilbe in large de gree for the Wider Size dis-

1

tribution recorded by idntzig. Unreduced pollen grains may

have been produced with greater freouency under teemperature

and 111ght cornditions to which hdntzirng' s material was subjected.

Mfintzing (1956) studied meiosisl in the tri oid from

Skane and observed it to be characterized by great irregularity

and the presence of trivalents, bivalents, and univalents at

metaphase I. The frequency of tri-, bi-, and univalents varied

greatly. Five to fourteen trivalents were observed in various

0. m. c's. The probable association of chromosomes in one case

was recorded as: 15 trivalents + 7 bivalents + 4 up1VQlents,

On the assumption that this interpretatioxi was correct, the to-

tal number of chromosomes would be 57.

First and seccond an.aphases were semetires observed to

be quite regular and other times rather abnormal. Lagging

univalents at anapbase I sometimes demonstrated a strong ten-

dency to divide. One p.m.c. at anspbase II showed 5 different

anaphase groups and a number of eliminated chronosomes. In

consequence of such irregularity tetrsds were often irregular

and polyspor g was common. Of special si gnificznc was the

observation that lagging and undisjoined chromosome pairs at

l l- , ”--~ \Chrome--aceet1:— formalin fi- idn Mltfl srort preparatory t*eat-

ment in CarnOv, -55—

anaihase I sowetimes connected the interphase nuclei, resulting

[,4-

in fusion of the second divis on spindles with consequent pro-

duction of dyads containing the full (5n 2 57) complement of

chromosomes.

Mdntzin; concluded that the triploid studied was auto-

triploid on the basis of the high frequency of trivalents at

meiosis. In some cases observed, 42 of the 57 chromosomes (74%)

occurred in trivalent associations. The autotriploidy conclusion

was also supported by the observation that one conspicuously

'large chromosome was present twice in the diploid and three times

in the triploid. he suggested that the clone probably originated

from the union of one reduced and one unreduced gamete from

diploid parents.

The author sugtested that diploid x triploid crosses might

conceivably yitld tetraploid (4n = 76) individuals. This cross

was later attempted (Kilsson-Ehle, lfiSb-a; Bergstrom, 1040;

Johnsson, 1940) with success. (See p. a .)

Study of meiosis in the original triplcid from Shane,

as tell as in several clones from other parts of Sweden, led

Johnsson (1940) to concur with the findings reported by hfintzing.

P. trenula L. diploid X P. tremila L. triploid

Cytological stulies of these hybrids, reported by both

Bergstrom (1940) and Johnsson (1940), were limited to root-

tip material since immediate interest was centered on chromosome.

numbers of the hybrids. In accord with the results'anticipsted

lSee pp. 25-24 for further amplification of this cross.

-59-

by Euli'tntzing the hybrids ranfled from the 2n n11.1Lcr of so to the

‘

.raploid number of 76. Tne rsjoritv were aneuploids but onewpvd

-

exactly tetraploid indiVidual was observed.

The difficulty of countingc10poover, 331ecially 1ar

numbers, in such material was mentioned by toth authors. It

was noted that counts were probably in erior on an average of

+ l chromosome. The Nile ipal difficulties involved were at-

tributed to crowding of small chromosomes or median corstric—

tion: of some chromosomes which may be ion:‘0

U

and uncolored,

resulting in 1 chromosome being counted as two.

Both worlmrs reported use of diluted chrome-acetic-forma-

lin for fixation of root tips. La Cour BEE proved to be un-

satisfactory.

Johnsgon stated that fixations made in early morning or

on cold, rainy days yielded few divisions but well-defined

metaphase plates which WJPS suitable for c0unting. Fixations

made at noon on sunny days yielded 1umerous divisions, but

they were unclear and without distinct metaphase plat.L

(D

S.

POpulus trcmuIOides Kichaux

First reports of cytolorical investigations in thisx)

f\

species were made by Erlanson and Hermann (1937). Studies

3 in p.m.c's. of both a strictly nonos exual Speccimen

and a bisexual individual were described.

Observation of metaphase I in the normal dioecious tree

revealed 18 pairs of autosom1s and one pair of heterochromo-

somes. Chromosome morphology was reported similar to that

‘00

ion: CQPHOV'S fluid or Flemming's; sectioned 12-15 u

: ironalum-haeuatorvlii.

-40-

described in 2. trernla n. diploid by Blackburn a Har~ison

(|924). The assumed sex pair were claimed to be easily and

readily distinguishable by theass authors. Both divisions

were observed to function regularly, except that sometimes

one or two pairs of autosomes lagged behind the others at

both anaphases. Normal tetr (i of pollen rains were noted.is.)

C)

‘

Studies of meiosis in tea n.m.c'3. of perfect flower

of the berMannrolite plant revealed striking and interes tin“

abnormalitiees of c1 ’3

Omosome behaviour. In the first place,

3 W93 TBCOPGGd in the order with which

the different anthers underwent reduction as contrasted with

the order and sm1m1roniza1fon in no mal male catkins.

Anaphase I showed, in some cases, very Ureat irret'w1la ’S itv

a

The KY pair (appa.rentlv undifferentiated morph0101ioally fromU

normal plants) sometimes behaved normally, With one member

going to each pole. Sometimes, however, the entire pair was

observed at one of the poles while the autosomes were on the

equator in early anaphase. In some instances, at this phase,

one or two pai°s of crromosores were to be found in the cyto-

plasm, having failed to reach the spindle. Lagging of the sex

chromosomes was not observed altho this, according to 1 e1n1nan

(1925) and Erlanson & hermann, is characteristic of thy dioe-

cious plants.

Counts made in interkinesis nuclei by Erlanson & Hermann

demonstrated the irrepularity of distribution of the chromo-

somes in division I. These nuclei contained anywhere from 7

to 21 chromosomees with occasional chrOFOsores lying outside

the nuclear membrane.

Division II in some cases on‘ccccad re'wl~rlv but sone

Ff”

Ho

('3

U)

’—

93

i

P 9 ,0

3

3

(I?H F

.3

-a-

:5 I)

3‘

-CD

CD

0 O 3 0)

:..J.

O 11'

.J

r.l

H

:1

§ 1 H

. ._ 3 ., . .. .w 1, ‘. ._.-- ,‘, °

ertra soincles were 909% 01 V'ipu tue c:~<*oso “s Wuic

(’1'

"1 a: (7)

\AJ

,3

U]

’L

U)

2":

CO

3 h)

:1"

S

}_Jc

) (a

._'J.

1D

‘4

Ho

.’4

C]

:,.

O H l~—1

H:

)0

1—Jo

H

D

24

.3 ...J

(at.

O

I

'3

l-_|

Th. chrowosomcs in t};se crses became distorted ani clnwoed

e O (D

‘3‘

(D

"S

L—a

OJ .J

4

'I

v-4

5-

'_Jo

l l-x’

U .1 J

H TD

H

d

1-»

(.0

{'1

("'1

L')

1’1

1...

C)

".

{—4.

l‘J 1

r 31

:c,

(NJ

1J

O "c

O 1"" A Q‘

I

srrniv iris r 01:orflv (W scrfivefi.- U .

A U)

’24

,4

p.10

1;:

1t be anticipated, the pollen studies made by tn

“*3

VIC? ers from perfectL \J

averaged about 37.5u in diswetcr. Out of a render sawp e of

50 grains, the sizes TPWQOG from 72.5g to $2.5p. The

from norr-ial 111.219 a span w- s rarltedl :7 {smaller 1 o di T‘l’flt‘f}1 av-

eraging elucit 97.ep. I arf QFQlHS ” re recorrled rarely if a

all in this pallen.

Peto (193b) 9130 rcnorted stn‘ies of feicsisl in p. m.c

of normal dioecious ralc flowers of P. tronulc fies hicrsux,

l _, 1‘ .1 q, _ _ -_. g _‘ _ o a, ‘-

_Lomrams exxiueu; '0‘0‘1. unziir1jLorl. szCé Lia

( '2‘!)

k) \I

1".“.1——A

'0L).

De mlfiT‘ZIOPMQtlJIl of lfll:iv‘lxn.ts st rmat:otssa:1 1w.s rerxnited

‘was rtd;(fliserved.

p-

f

.- .- .‘. ._ “- ,._ 2., v. r V. D I. V'- - .. " ,_ '1 , _.‘_ ,1,

11 1! gOW‘JJHLd»€3 Oi wifarwn:nlj‘£ oor,,3011111 HWC iuitcn

-* w #4» » 2- ~ ~.: -1 . fir. . ,

both Of tic c (c1 ens stucied hr 32,0 11; one ._ovc€ a very

Wide range in collar diameter as 00““vrcd to the Grier. One

- ... _ " J '7‘: _ ‘l ‘. ‘1 _ .. ._ - . : _"_ _ , - -’.r:y

Kwarie1111ctx£wsi 19.Lh1 QHCL1M3.FW’ (Hclxfiftek ”ffnl 0:11 QLLI*- Q0,

~." 3 ‘t r" _' ' 1, ', I () r' a _

ocpc: hwy LL oesween no.4n “SC

“'7" / ‘ — ‘ v 0 H '1

i;ixatior- czl kawrvt tn alcOIOi, gihcial acetic nniq) and

- .. r7 , . 2 .- L ,° - - 1-. 1 - ,.storave 1? (CW filDOio], sue ‘ 111 Iludw; C ”in: acgto—

O

:ic bivalert reported bv Erlsns On 3 Hernann

18

. ‘ "“ t r s '\ ' . n ' ‘ ‘ '

VCVGVVJ mitN iw‘ 37.5U “‘pfl°tflfl J E l” “A” 3 ‘KJ Hun, Peto

" -" "— ~ J a L 1 u " f ‘H .1 V F‘0 3 . :‘V . ' o I " \

Sjluv'fger‘ 1‘ frp L‘ QC V 1 - .Iq ‘ 13-1 L‘ 1 . ‘ T 0 “081-3988

fiJ‘ l P n” Ju“ 41": 11/3 ru‘b’.71f\‘n(‘c‘ “If“ YW‘W'hpy- r. 11p n'3*f“v'~()‘n r.) lor‘ 0‘1:1. L. 0’” C) 4 K’J C K‘ - 1C . u - .. " )|)(. i v‘ --’--—"'. ’ J O - .. V ';.‘4.-‘_ U...‘ a . L - k.' VI ;)L_'L)-

-. .9 V} . J ‘ w r79" r15: '3 _‘_ - u 7 , v .

tested tsws (Uorw1 tLvu9niL 6:8 01 :£-;s coprsus E. pUsVUil-

i v‘ I“ 4. Yl +""l(—\ 1:k‘OT1'1-5_O 14*“ VJ} .LJI rig}, v.‘,r_3Li \3njm 1’ 0(313-01 .7 (a. (3 ( 3. w If.“ (3‘. C‘ V, n) (‘IYW

L‘v —'<“ ml" —t -‘ » ~ U - . _ ,. » -. . _. .__ _._n V v. _, 1‘ .. _.. s I- N _. ._ -N

v-° . . ~ .. ‘7 -V ‘ . . -- -"- ~ -' , ,, ~ . 1- -~ J. ' .91 '

CU t 1V“) wag Lflvc Dewn U CU.{RlbL{lHJ fswtor to 5N1? UtLoi-

.1 .- . .A‘ .. -- -l, ,. "‘ - V,

A £4W’99M: 1 Kris 1:o:wnsl iji timeunntx3Ui91_ 51w:¢ivséizlui.1zne

suthor belie v.9d th:t dUads are likely formed thru fsiluve of

'...'0

CXtOfihes's 9rd micros, no: - 3911 formation 3

W911 formation did not 'lrnvn mommy 5w'1

-1

.aJ

( ,.tzat CEtOJlNBSlS 9

- n ‘ _ .L. .. . “a. I“ j ‘- «- r‘ E 'V

the interphaso between 6ivi31's I ans II. Tnegu oncesses Hose

frequefif317(fl3S€PV6d.41317; 6619*9d rwflfljieafter telxnw9se of the

second divi. Hiir. Peto cor:chLc<d tht t WQY, under cert9in

1.1-. ‘ .4 ;I+_ _o ‘ ‘0 ‘a 1 '1

PCWWIlLlflflqsy I»:r~JJJ 33:1,1t . Jon r11019i. to Lye anfineo.‘

If this tonooioy to produce diploid grwwns ozn be indnoed

it su rests an excellent prortunity to produce triploid or

tetrsylcid plants. Tls Untursl occurrence of auto1iloid

olents of P. trenula in Sweden 1d P. slbn 9nd P. Qfifi‘QOCPS

in Canada provides ovidefise ,F9t diploid pollen of o‘be“ species

in the gynnis is not IHKMTW“0K 3111M3tUPO.

POffifibJS Tflfi._d.’-CU<ITtnta hfiichawu;

POtO (19:75) PCPOT'tCC; )5?!“sz 1'", cytolokjicgl 811111163 in tfiis

elded 9lwont id9nti391 results 9s {?Cse observed in:13

J (D

K)

*JO

'3 :9

M

"Petxa ditixiot, ?vvno\mmé, 991Kt9t 9(fln191117 seeixyj 639173 Wifiiti

p.w.c's.

| g.

-44-

tre tvo P *“o*Wloir€s Cfvciwcns inve ti flied. POllen nnfily‘. -_,.

3318, however, showed c Wider Variation in foiled dia*eter, i.c.,

17-5“ to 49.7r (weiLfilxriirwwn fiewoter 2 93.01),

Porn 11': ol‘no L,

’fiottstein (l?S?-s) reported :, alts u. a” o ndcw l diploi

wit? 19 pairs of chro oso es but Peto ( T3£) studied tfio speci—

rcns and founc tram totn triglo C. Lil?nwijn (lEZ§-s, 1939-b

10/0) 2150 reported stud; of a specimen and rego ted it as the

first triploid POuulus found in the Eetberlsrds. Feto (1938)

used ? trird representsiiye of th’s sLecies in a cross (P. 9153 L.

X P. i2"flfliid9flt9fflluiChQUY) BJAlZPepOPtEIiiflTlt all of’tlx:1»fbrids

exeminec were exactly diploid. he concluded therefore thwt this

particular individual was diploid. From the cooperatively meagre

information availsolc it would 9930 r tint the diwloid form does

exist in nature, and protatly in greater number than the triploid.

The two specimens studied by Peto were leveled Al enr A

Al 3 P. elbaz A2 3 P. Elba var. nives snreo-intertcyte ?. Tne

mean frequency of bivalents enc trivalents at we sphese I,

based on a study of several nuclei of each specimen, s:owcd

the maximum (i.e., lei-J, the haploid nmiflf'er ) configirations

present. A9 however, snowed a: everoce of 13.85 trivalents,

whereas in Al only an avers e of 5.8 trivalents were observed.

Peto concluded thst A2 was an autotriploid and that Al may be

Percentage of apparently good pollen was hirh in both

Al and A9, vein” )3 and van. Furthermore, dis fiibution of

pollen sizes was narrow, with no evidence of urrcroceo pollen

grains oein; pron100d.- Actual size dis

cided quite (310331 with those resorCeC

fcw' irdgjloici f} thfififlllfl 14.:

F. 91,0.” 3 5301' to 711411.

E. tre wig :‘esg 10'421

logically ant1c H.

B

tins CCH18:71 otcnis rmsuo+th 3;?P6k:l__dfiitjxfis

end trernld trigloids 1.ler1 snow lov 1n

pollen and a amnvvfil siZe distribution.

exrflxnwl tiCfil fov'ifl is e11uireot jJUZOHSlfio

bslsnced pollen grmirzs prob-“11.1131 111:; >111

to such an extent th.t it i ir1os1:itle

fiverflieirzy'fiwt Ivoliler I sir I'Lt".COILseC1.Le

that degenerate 1.1101; slow 1 and could L

e. h‘lfitfl‘i‘T‘JClWB lJlCllLdEll 111 "(Nird's 1)? 17

Fermi 1 slim L. di1‘iloid x 1:. (Lucident

ordjnititfli linfiqts Cffirr-

by Tonetorp (1957)

O

t3 't plrnwts 1” owdjx

5:10:31rqiC1 1y alts

icean_ of good

re.o s1;;~sted an

1 e. n . ~encg. Genetically un-

ate vcrv rdnidly and

to determine their

ntly onl~ pollm

e recoQnized

oor pollen."

eta icne 1,1}:

- 1

Peto (1956) detETHl eo the someti

end S t1;died meiotic behavior of l2 of t

discovered near Ottawa, Cens1e, fi 1t r

r, r‘

C (“’01ILCSOZTK)\/ 3113311”)e r ( 5C5 )

hese natural hybrids

>>orted by Feiu1ur3er

(1930, 1940). An avereée of J bivsle were observed et

metephase I in 10 of the trees. 111sr~ta fre"uenc ies tore

not determined but the proyortion of ring to rod bivalents in

thes lO hybrids similar

grandidentata h Peto titereiore

degree of homology meteen elk 0(.1.

‘1‘":3

4s).C::Z"OI:‘OS

demonstrated in

In one, lo. 2 unimv

- A. I?

to tnst noted in P.

concluded that a high

the other two hybrids

lClrtS l().9 1)_slui

Vere observed, while tlv other Slowed 54.4 univelents and onl

1.8 bivalen 3. Some nuclei in the latter showed complete asynup-

sis. Peto believed that the asynapsis oew‘mstrsted by these

two plants was not due to low homolorv betvern tn1:?

parental

crromosoJmes or to unfavorable environment, but was an example

of genetic factors limiting pairing.

Only‘iwna triV’alxxits werwa<fi3servemi in alflL the all“? x

n

crandideutota nuclei examined and Peto therefore concluded.LL

tb’t structirsl char; 1s have slaved an imoortant role in the.L L' J.

evolutionary differentiation of tbese species.

Of special interes1 is file observation that tie plant

showine highest degree of aSjrapsis nevertheless produced not

tie standard devietionr—«J

apparently normal pollen. In ed<ition,

of pollen diameter was relatively low. But amen r the hybrids\J

'I

whicn exbitited normal meiotic pairing, wide variations were

observed in the perccentage 01 good pollen, mean diemveter, and

standard deviation.

Eleven of the trees studied by Peto were typically male

but one was apnarently oolrvtmo-ronoeCWQus with both steminate,

oistillate, and, presumably, perfect flowers.

P. elba L. diploid x P. tremuloides Kicbauxl

Three natural bybrids of this cross were studied by Peto

(1978). The somatic number of 5% chromosomes was observed.

Similar results with reerd to meiotic behavior at meta ptase I

were observed for the three trees: 16.4 to 15.8 bivalents beiro

observed. lean size of pollen corresponded quite closely with

1These ryirids were also discovered near Ot1amva and reported

by Iieir‘rfourger (123:40) .

some of the al‘na x Jmcdidentata l'ybrids. One of He trees

has nonoecious.

On the whole, these Tvtrids exhibited less vigor than

the alba x nrandidentata hybrids and Old not Lave as promisingA.)

UOSSibilitieS.I.I

Popnlns cwvrescens (Rita) inn.

This snecies, occurring in both the dicloid and triploid

forn, is believed by Peimburrer (1950), Wettstein (1937-C),

1 a 1 O 'Vw *" v' 1

and others to we a ratursl hybrid: P. sins L. X P. tremUla n.

Peto (1956) reported ciromosome numbers in 4 specinens, find-

ing two to be diploid and two triploid. studies of reiosis

were reported only for the triploids.

In one triploid 18 Divalent and trivalent associations

..

were observed, whici approaches the theoretical maximum of 19.;

1

The trivalent associations were in use minority and the author

G)

concluded this to be evidence of hybridity. The 0th r triploid

cerescens exhibited on the average only 2.7 bivalents per nu—

cleus with a range of O to 7 and no trivalents. Univalents

were observed which divided at the first division. The author

concluded that lack of homolosy among the three sets was an

unlikely cause for this high degree of asynapsis and that gene-

tic factors limiting pairing might be present.

Abnormalities of meiotic behavior are reflected in the

wide variation of pollen diameter reported (19.nt to 41.8u)

which appears to be bimodal. The author suspected the presence

of unreduced or triploid pollen grain . Percentage of presumably03

lPeto (1938) stfltEG that 7. canescens is very similar in appear-

ance to the natural hybrii

differir°

as of the cross alba X grandidentsta,

U only in leaf shape.

- A k“; -

good pollen, on the other hand, was found to be surprisingly

higdl (9htf). ’Thixa cormriticr1zis pumflmnblg'eytrlaijnible (M1 the inureA.

basis as that su55ested by Pete for P. alts L. trinloids, 1.9.,

that non-viable pollen dis 1rterates rapidly and is not reco<5niz-

1‘3

ble as mnfliat maturity.

o -. o ’ -. I q- . ~, 1 - 1

X Populus canadenSis huipnei (bimon-nouis) acnelle

Studies of p.m.c's. in this hybrid were reported by Peto

(1958). The somatic number of crromosomes was found to be 58.

neiosis was extremely irregular with urivalents vauying from

O to 10 at netaPhase I. The irweui'rjtic of meiosis were re-

flected in the pollen characters. There was found to be only

I

1 1

GW good pollen and the sca1dard aetixiion of pollen diameter

was the larest oi any of the several species reported by Pete.

The author concluded that the hybrid constitution of this-

plant accounts for its irregularity.

Popplus nigra L.

Blackburn & Harrison (1924) indicated that their descrip-

tion of clromosome number and morphOIOgy in P. tremula applied

as well to 3. Lifira (see p. 54).

Dillewijn (1940) on the basis of secondary association

observed at metaphase I in this species and especially in the

variety it-lica concluded the 5enus Populus was a secondary

A -(‘

l—Jo

+ {.7

9.3

S O W H

3

3

I...)

C D (D

H

mly‘loii nurber of 8. Of these, 4'

chromosomes have presunably been cz=ipljc atgd and 4 duplicated.

lConsidered b? Telder (1940) a variety (o01

from the cross P. deltoides x nipra = X P.

but believed by_Menry(lul4),'EFfiHEider Tlffid , and houtzag-

ers (1957) to be a hybrid from the cross: P. re:Heiepgta

Henry x P. nimra var. italics. 11_ue:1..chh.'- _. _...._.\.-’.._.__ _..___..-__._..

form) of a hybrid

con-ec ensis Inloench.

-ao-

‘1

f.)

F-

e prescrt number (1%) is believed by tlis worker to

n 1 . . :- ‘ ‘ " ‘ "I1 "" . i . 'r” n. J‘.’ " :"‘ ’ ‘ ’ '

l tusior oi d CLIOWOSO es hltdln one o; tLG .P\LPS of"3

d)

U)

)2:

}_J

C—f.

0

three. Division I and II were resorted normal for this Species

as well as trxa‘yflrieir italicaq Ixillewiin obseiunxi thtt the

rrzclfiwdlus iii P. rd_rra :fi:owcm1i3 or'rnore trids c

3 q u ‘ 1 _ _ ' . .‘ . ' A , . . v

Pcnrilils (39117031193 'VHI‘. 1L1;r;odqfiwetnris 1.8Di‘fW

(W ,‘

Accord5ng to Rehder (qu0) this is tn: 3. be

bslsomiferfiL." and also of 3. tricronqrgg Hook. Fe reportedly

concurred with the 5eneral morphological descriptioq of cnromo-

somes for the genus as noted by Vlnctflurn & Harnison as a result

of toeir invesliqeticn of E. trvfinlw L. and E. niyra L. T~"e did

\ J

I' - — ." .—-.~. ~ —. -- . ‘-‘ "“‘\ rv "\ 1" ')‘,' ’ I - - ‘ '1" " 1! ..‘ ,‘

ncdl, ledBVffl”, alfree .ritzr tne;»» scu -GIT3' I¥?PQIW, oi tyma so; {tic

u.

Crrowosone number of 76. leurman reported this species and m

trichocaroa Hook. as rormal diploids, 2n = 55 Heterociromosomes

were observed ir both spooles, so that in

or 15 § V, and presumabl;'ranloid fewrle 3 lb + X.

~,v— .,- 7...- ”fir: 1: n .,° .0 . . .

Lille iyn's (I940) studies 0; mulTSiS in this spa01es

: 1" -‘ _. 1‘ ‘ W” r~ ‘ ‘ _' _: . .0 _: . ‘7 ““1 _ O __O '1 _ .. O

(variety) reVealed flu'eflnkfl «gictic ir:05nlarltigs. MOD-pulp—

ing and lQfiTIW" of ctrowoso es resulted in tne nresenco of extre~

nuclear clromatin ard swindle fusion st division II. Pollen

.1

dicru tor SEWrmxi Hide vnrdx“tion asscmfixvted riftvruwrv cases Cd‘

triad, ékyui, and worsd forwaiio-. lfl:\wss concluded that the

IQNTBTWNIG id??? fillfildjtifm3 CCnifiIand 11§e \fiier/lielxi hit seine swig: ad- _. .

U

‘n (l?iu) consisted of: fixation in

i 7 hieroform) or CCPNOV s PE

‘.-. an.‘ ‘. ' 1

(31.1 IJH‘E) .1-.“ 1,0" ”POI/€10 t0

I L.

"‘

_

j1';

QA

5D

‘4

Cf

-.-J

(D

r1

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er

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that this variety is s hy’orid. Jillev-Iiin found the Theoloi Ll.

number to be 19.

(l) X P. brsbentics HOUtZUSGPS (n = 19)

__i-

nlnnteriens‘s

(2) X P. Nelrics houtzn ers (n = l?)—- 1‘ . J

(5) X P. PODlst? Scrneid. ( = P. ijVlatg flit- X 3- H1C“97‘- --fl (“4? V . fl ‘ 'Iv“’v‘| "T— 7 T "' x(s wkuninisT so -+ Ll., Val.) (. _ l.)

' -\r‘-¢‘ >1)1Hrw°n m '1 ‘1‘. ("r J".-

LL L; A 11’1(4(<-'C3j--T n)?’ 0 11.119; 1 '1 Jr. -1-‘.. ‘ U

. ' if) J~" — .r'Y- , ~ “"71101*tef1 l“. {is elm: l'QLdlgf%fl INITJJD51 of?

.J

(F

L O 1")

e {Jo

O U)

l.Jo

Ts generolly 3330-

. - '1 --_I _v_1_ a, "-11.. _ : ,3 .0 -.. 1.. 1‘ .5 _2 _ _ __ 1 _ _‘

ciatet hl-U tuuULULLJ were nOLed in these plants. BuldinQ

of the nucleolus at prepnsse was observed in all. Pairing

was good in (3) but poor in the others. Polysoory was cnarsc~

tEITiS+i'3 of‘ all {N“d 11h"”l*" cbrwr oscw cs pinns euvtraryuzleais.. , ~)’

rrotcin was ooserveo in (l) and (4).

5H

Dilletijn (1339~b) concluded that tte constart broloida.

~ .- \ (N . L ‘ 1 -\ - ’w ‘ ' ’ ‘* 1 1 ‘ I" "‘ ‘ ' ‘ \ .. r— - ' ‘

Inniben? (14%) 1.. th 1Jenru: P.1'hl_m. arul KIN? frmwiuern. fornmatitm:

3‘ ° . ... . .. . 9 ,~ _ -.. . 1—. , - '

of dyads in many species are factors favor ole for brewd1n5

work. He concluded, also, trot tEis probably eXplains the

-1 o .. . -~. - _ ., ,\ _ a -1 _ ._

polyxor.nism #11 in the terns an: tne occqronce of naturulr «

triploids.

CYTOLOQY CF SEX IN RNPVLUS

"e presence of

bet nociromosores in use various species reported here but no

effort has been made to emobasize this aspect of the cytolo-

o ,,_. __1__ _ '1 _ , V .r_ f _- . 0 l __ -0 ‘I .,, ._ .11 r.‘ .

rfiLcal_ moiha. I11 at..it AN; cc) t_M3 CiLm4fil‘d‘S nu‘evjxdusiuy rewie 419v A

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--------- (lCl4-c) A new ‘yorii poplar (foculus nenerosa).

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""""""""' (1938-13) (The glant L30{D)-Ei‘f"--tile recent swed—

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-.‘37-

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stungss tei:gerung

1a. (lncreasinL

t “Oenance in

34-43 (Cited

Chin

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------------------------------ (1939) Die Jermehrung und

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()\:_/

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1933<

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Al3eneen lieu113sdienst (H.A.n.) Pp; 20.

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1 V

1

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a s

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Y

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CD

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~59-

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Jap. Jour. Genet. 12 (2):

-50-

f‘r)()’— 7(_; ,

numbers in some An5iosperms.

101—103.

SALIX L. (Willow)

The willows, because of numerous and often polymorphic

species, are, taxonomically, one of the most difficult of

tree genera. In addition, many of the species apparently

hybridize freely in nature, thus contributing to the general

confusion. About 165 Species are recognized. More than 65

species are native to North America but only about one-third

of these attain tree size (Rehder; Harlow & Harrar).

Salix nigra Marsh. of North America and §, glbg_L. and

‘§. fragilis L. of EurOpe produce the bulk of commercial willow

timber which is largely used for artificial limbs, boxes, crates,

woodenware and novelties (Brown and Panshin, 1940). A number

of others, both trees and shrubs, are extensively used in the

basket industry or as ornamentals.

As a group, the willows require considerable moisture

and in consequence characteristically inhabit stream and river-

banks. It has been said (Holman & Robbins, 1954) that in

this genus the guard cells have lost their power of regulat-

ing the stomata and the latter remain permanently Open. It

may be assumed that this feature restricts the willows to

habitats where there is abundant soil moisture. Investigation

into the cause of this phenomenon might prove of genetic

interest.

ARTIFICIAL SELECTION FROM WILD STOCK

Record of conscientious effort to select superior indi-

viduals from wild tree members of this genus have not come to

the author's attention. There is, however, abundant evidence

that empirical selection methods have been in vogue for many

years. It should not be overlooked in this connection that

forests and stands under forest management may be subject to

a silvicultural sort of "mass selection" which may, depending

upon intensity and preper judgement, result in some improve-

ment. Individual selection, especially in EurOpe, has undoubt-

edly been carried on among the various tree and shrub Species

with their various varieties and natural hybrids which are now

cultivated for the basket industry and ornamental use.

Of special interest in this respect is the white willow

(Salix alba L.) of EurOpe and its several varieties. §, alba L.

is widely planted as a timber tree; §. alba calva G. F. W. Mey.

(in the literature usually: ‘§. alba var. coerulea Koch), the

so-called "cricketbat willow"1, is reportedly superior for the

use which its name implies; §, alba vitellina (L.) Stokes, a

variety used for basket making;‘§. alba tristis Gaud., var. is

valued as an ornamental. Still other varieties and forms of

this species have been described. In addition, the following

natural hybrids, presumed on a morphological basis by taxon-

omists, appear in the literature (Rehder, 1940). Since def-

inite experimental evidence is lacking as to parentage it is

impossible to indicate whether §ali§_glbg L. was involved as

the male or female parent.

1

Henry (1914) recognized the cricketbat willow as a hybrid

between §. alba L. and E, fragilis L. Schreiner (1957) indi-

cated it as a hybrid (X §. coerulea) betweenl§. alba L. and

S. gracilis. On the bas s of cytological examination, Wilkin-

Ebn concluded it was a variety of §. alba L. and not a

hybrid.

Parents1 Name 2£_hybrid

S. nigra Marsh. x §, alba L. X'S. Hackensonii Dode.

S. fragilis L. x §, alba L. X'S. rubens Schrank.

>4

S. Ehrhartiana Sm.S. pentandra L. x §, alba L.

S. babylonica L. x S. alba L. X.§. sepulcralis Simonk.

Numerous subspecies, varieties, forms and natural hybrids,

involving other species of Sglix are described in the litera-

ture. The more important and more widely cultivated of these

are enumerated and described in the work of Rehder (1940).

Johnson (1959) lists 42 natural and artifical hybrids in the

genus.

CONTROLLED BREEDING AND HYBRIDIZATION

The first controlled breeding experiments in the willows

were conducted by Max Ernest Wichura between 1852 and 1858 in

Germany (Wichura, 1865). A n accurate and precise worker,

Wichura's interests were not disconcerted by the "practical

possibilities" but were directed toward a contribution to the

rapidly accumulating genetic data of his day.

According to the theory of the time, a "species" was con-

sidered to be an integral whole. In Wichura's concept, there-

fore, a species crossed in its entirety and not as a number of

characters taken as units. In all Wichura made thirty-five

successful "binary" (interspecific) crosses between twenty-

one different species of willows. He reported a much larger

number as unsuccessful (Roberts, 1929).

l

Cytological studies (Harrison, 1922; Blackburn & Harrison, 1924)

indicated all parents except S . babylonica L. (which has not

been studied) to be tetraploid'(4n =f76).

-6 5...

Wichura noted in his willow hybrids, as indeed others1

had in other plants, that hybrids demonstrated greater vegeta-

tive vigor and were less fertile if derived from crosses be-

tween species of distant specific relationships. He also veri-

fied GHrtner's observation of the identity of reciprocal crosses,

and, according to Roberts (1929), the following is the first

recorded, complete categorical statement in the literature of

plant breeding with regard to the function of the sex cells

in amphimixis:

"From this (identity of reciprocal crosses) 1t.EE§E

follow,however, with mathematical necessity, that

the ppllen 0611 must have exactly the same share in

the conformation of the fertiliZation product as the

egg." (Wichura 1865, p. 86. )

Wichura's concept that species crossed in their entirety

undoubtedly confused his analysis of F2 generations. Perhaps

he did not grow sufficiently large F2 pepulations. In any

event his general conclusions indicated that segregation did

not take place and as might be anticipated the post-Mendelian

workers made haste to disprove him, at least with regard to

special characters (Sirks, 1915; Ikeno, 1916, 1918, 1922;

Nilsson, 1918; Lotsy, 1920).

It is worth noting that a pre-Mendelian American sali-

cologist, Glatfelter (1894, 1895, 1896), recognized in wild

populations of Salix nigra Marsh. and‘S. amygdaloides Anders.,

intermediate forms of great complexity ranging from one parental

extreme to the other. These variations he attributed to hybri-

dization between the two species, crosses between the hybrids,

and between hybrids and the parental forms.

I»

Kblreuter, Gartner

-54-

Some directed effort in breeding willows has been stimu-

lated by demands of the basket willow growers. Schmidt (1957)

attributed the unsatisfactory state of the product in Germany

to the impurity of the material available to the grower. This

condition has been brought about through natural crossing

among the cultivated forms. He recommended that the initial

step in breeding basket willows, as with other economic crops,

should be the production of pure lines.

Wettstein (1951) reported initial breeding steps with

Salix americana Hort. (possibly S. pgrpurea pendulg Dipp.).

This form, valued for fine basket-work and as an ornamental,

is characterized also by a bark which peels with ease. By

cross-breeding it is hoped to increase the tannin in the bark

which, in turn, would not only act as a protection against

animal pests, but also provide another product. The Fl's from

crosses with S. caprea viminalis (?) and S. purpurea L. are

said to be promising and will be bred further.

Barnes (1951) reported insect resistance breeding work

with several willow species. In addition to the hybrid

‘S. viminali§_x purpurea (S. rubra Huds.), formerly found to be

immune to button gall (Rhabdophaga heterobia), 4 other va-

rieties were isolated. In addition, immunity to attack of this

insect was found inl§. viminalis L.,‘§. purpurea L., and.§.

.ElEE L. Further breeding work is planned to produce new va-

rieties resistant to other insect pests. The physical or chemi-

cal property possessed by immune or resistant forms has not been

defined.

-65...

HYBRIDIZING TECHNIQUE

Members of the genus Salix are normally dioecious. It

has, however, been known for many years that monoecious plants

do occur among individuals of certain Species (Shimek, 1895;

Chamberlain, 1897; Burkill, 1898; Engler & Gilg, 1924). Some

idea of the frequency with which this condition may find ex-

pression is indicated in the observations on‘§. amygdaloides

Anders. reported by Schaffner (1919). Of 100 Specimens of this

species which were carefully checked, 9 intermediates were found.

It is of Special interest to note that aments with staminate

flowers had these flowers limited to the lower part of the

axis, while the carpellate flowers were found at the tip. In

addition there was a transition zone between these two areas

which seemed to be neutral in regard to sex. Bisporangiate

flowers were frequently present as were also abnormal flowers,

i.e., flowers with structures develOped which were partly stami-

nate and partly carpellate. Schaffner believed sexuality in

this Species to be traceable to physiological conditions.

The occurrence of monoecious (or even polygamo-monoecious)

individuals in this genus is a point of some technical import-

ance to breeders. Aside from the apparent advantage of effect-

ing artificial cross-pollination in dioecious and monoecious

plants, the monoecious condition permits of true selfing. The

only drawback in such instances is the possibility that self-

incompatibility may exist. According to East (1940), however,

the monoecious plants of this genus appear to be self-fertile.

Sex and its modification in this genus is discussed in

more detail under "cytological studies," p.‘10.

Yanchevsky (1904) is reportedly the first worker to

demonstrate the method of using cut flowering branches of

.§El$§ which produced mature seed in the greenhouse. Wettstein

(1929) reported a number of successful crosses between various

willow Species using this method. He emphasized that the flower-

ing branches Should be cut when the first swelling of the buds

occurs in early spring and placed in water in the greenhouse.

Following the appearance of the adventitious roots the branches

should be transferred to a nutrient solution. The cuttings

should never be placed directly into soil or a nutrient solu-

tion as vegetative activity is emphasized and the catkins

usually fall.

This technique may be successfully used for crosses in

the open with species which do not mature their flowers at the

Same time. By controlling the temperature, cut branches of the

male parent may be induced to produce mature pollen coincident

with the receptivity of the female flowers.

POLLEN LONGEVITY

Nohara (1924), a Japanese worker, reported longevity

studies of the pollen of a number of species and hybrids in

Pollen storage was conducted under various combinations

of conditions but most successfully (i.e., for 70 days) in a

desiccator in a dark room at a temperature varying between

80 - 14° C. The exact percent of relative humidity was not

determined but he indicates this as the most critical factor and

that it should be low.

For germination studies pollen was germinated in a film of

2% cane sugar solution on micrOSCOpe slides. The slides, after

sowing with pollen, were tht in a moist chamber at room temper-

ature. Pollen was also successfully germinated in water and in

solutions of grape sugar. 'gglix pollen apparently germinates

very rapidly. In some cases pollen tubes of considerable length

became visible after 50 minutes when sown under favorable con-

ditions.

Nohara's studies led him to conclude.that there was no

difference in germinating or fertilizing power of forced or

naturally matured pollen. In his work with preserved pollen

he found that as long as pollen was capable of germinating it

was also capable of fertilization.

Bogdanov1 (1955), a Russian, has also described experiments

on pollen storage in Sglgg. He first worked with fresh pollen,

determining for each Species under investigation the best condi-

tions for artificial germination. Following this, he determined

the normal germination percentage of the fresh pollen for each

Species. There was considerable variation observed between

species.

Pollen.was stored in a refrigerator at -5° to+5O C. He

reported that both excessive moisture and dessication reduced

germination and preservation in a dessicator was definitely un-

favorable. The percent of relative humidity in the refrigerator,

however, was not indicated. It may be assumed that storage was

in the dark.

‘1‘

Original work not consulted. Data from abstract in Imperial

Bureau 2: Plant Breeding and Genetics -- Plant Breedipg Abstracts;

VoI. 6, entry #651.

Pollen of g. viminalis, §, acutifolia, S. dasyclada,'§.

caprea, and the hybrid‘§.,phylicifolia x S, viminalis, stored

for a period of 20 days under these conditions, left normal

germination unimpaired. A certain percentage of the pollen in

each case germinated up to the fiftieth day.

In Bogdanov's original article the most suitable germi-

nating media and the duration of viability of the pollen are

tabulated for the above mentioned and a number of other forest

Species.

SEED TREATMENT

éfllli seeds are known (Holman & Robbins, 1954) to lose

their vitality rapidly and should therefore be planted immediatea

1y after they are discharged from the capsule. Wettstein (1929)

emphasized that they should be planted on the surface with suffi-

cient light and moisture available. Even the slightest covering

of the seeds should be avoided Since lack of adequate light in-

hibits germination. Willow seeds planted under proper conditions

were observed by Wettstein to have their cotyledons visible in

3 hours 0

VEGETATIVE REPRODUCTION

The willows are readily prepagated by cuttings or root

Sprouts. Various species readily strike root from wood several

years old (Toumey, 1916).

CYTOLOGICAL STUDIES

The monoploid or basic number of chromosomes in the genus

§21i§ appears to be 19. A polyploid series, based on a number

of lists of chromosome numbers and other reports (Harrison, 1922,

1926; Blackburn and Harrison, 1924; Meurman, 1925; Sinote, 192a;

Gaiser, 1926, 1930; Tischler, 1927; Maude, 1939) is suggested.

Diploid (19 pair, as'§. purpurea L.), tetraploid (58 pair, as

§, alba L.), and hexaploid (57 pair, as g. Andersoniana Sm.)

species have been reported. The reduced numbers 22 and 44 were

also observed by Blackburn and Harrison (1924) and Hakannsson

(1929) in some species and ca. 24 by SinotG (1929).

HETEROPLOIDY AND SEX

Blackburn (1926), Harrison (1924), Sinote (1929), and

Nakajima (1957) have reported a large number of diploid species

with an unequal pair of chromosomes. Several tetraploids and

at least one hexaploid (§, Andersoniana Sm.) show only one pair

of heterochromosomes. Harrison (1926) has, therefore, logically

concluded that simple chromosome doubling could not account for

the origin of polyploidy in these species, since in that event

one would expect to find the heterochromosomes doubled also.

Lindsay (1950) observed that the evidence of the presence

of only one unequal pair of chromosomes in tetraploid and hexa-

ploid willows conflicted with.similar observations in other

dioecious plants. Hagerup (1927), for instance, reported the

presence of an XY pair in the staminate plant of Empetrum

nigrum, a dioecious:3pecies, and two XY pairs in E, hermaphro-

ditum, a tetraploid hermaphroditic species. Lindsay concluded

that in Salix it appears that dioecism is not interfered with by

polyploidy, even though the XY pair remains unduplicated and the

-70-

balance between the sex chromosomes and autosomes is obviously

upset.

Blackburn and Harrison (1924) state with regard to their

observations:.."some evidence exists of heterochromosomes, pro-

bably sex-determining in their import." Nakajima (1957) con-

cluded: .."the unequal pair is assumed to be a sex chromosome

complex of the XY type."

From the literature at present available it does not appear

possible to define with any degree of certainty the means of

sex determination in the genus §gli§. The work of Schaffner

(1919, 1955) and Harrison (1924) indicate that sex reversal or

modification in this genus may operate apparently independent

of the assumed sex chromosomes. The basic fact, however, that

a consistent differentiation of sex among individuals of the vari-

ous species does occur would seem to indicate that an hereditary

mechanism is involved. That this mechanism, under the influence

of physiological or ecological conditions, can be changed does

not invalidate it.1

Hybrid Analysis

Nilsson's (1918) genetic study in Salix has previously been

cited as part of the post—Mendelian work criticizing Wichura's

earlier (1865) conclusions regarding F2 generations in the genus.

Eleven years later Hgkansson (1929) reported an extensive cyto-

logical study of the F1 and F2 generations of the principle

cross which Nilsson reported: Salix yiminalis L. x §, caprea L.

T

See also Yampolsky (1922), s1not6 (1929), and Smith (1940).

-71-

Hakansson found both parents to be normal diploids with 19

pair of chromosomes. This verifies the work of Harrison (1922),

Blackburn & Harrison (1922, 1924), and Meurman (1925). Harrison,

however, noted that while §. caprea L. is in the main a diploid

form, a tetraploid race indistinguishable in the field from the

commoner diploid type was found.

Meiosis in the F1 hybrids studied by Hakansson was found to

be regular, except that at metaphase II irregularities sometimes

occurred which resulted in formation of diploid pollen grains.

Most of the F2 individuals had the diploid number of chromo-

somes and showed more or less regular meiosis. There were, how-

ever, two polyploids. One of these, a male shrub with giant

leaves was triploid (5n = 57). The other was hypertetraploid

(4n 8 82 - a4). *

In the triploid at metaphase I, 17 to 19 trivalents were.

observed. When less than 19 trivalents were observed, the others

were each represented by a bivalent and univalent. Some pollen

grains with the unreduced number were found. The pollen was

rather good, about 40% being normal in appearance, though their

size varied. Hgkansson suggested that the plant arose from the.

fusion of an haploid F1 egg cell and a diploid F1 sperm, the

latter being the result of a chromosome doubling in division II

as previously observed.

In the hypertetraploid at metaphase I of the embryosac

mother cell there were observed some univalents (6 - 8) and about

58 bivalents. Quadrivalents, and more often trivalents, were

sometimes present. Anaphase I was extremely irregular and the

plant was completely sterile. This plant was believed by Nilsson

..."70-

(1918, 1955) and Hgkansson (1929) to represent a synthesized

.§° laurina (?). Rehder (1940) indicated the parentage of §.

laurina Sm. as: §, caprea L. x'§. phylicifolia L.

Nilsson (1951) reported later duplication of his original

cross.§. viminalis x §, caprea L. and in the F2 found what he

described as an undoubted‘§. cinerea L. plant. This Species is

reported a tetraploid (4n ' 76) by Harrison (1926). The

cinerea-like plant's chromosome complement was not definitely

ascertained by Nilsson, but it is said to be at least triploid

and in all probability tetraploid (amphidiploid). It crosses

readily with é. cinerea and is completely sterile with both its

diploid parents. It is regarded as an experimental synthesis

of.§. cinerea L. by Nilsson.

Hakansson (1955) reported cytological analysis of several

other species and hybrids between them, including: '§. nigricans (?),

(§. myrsinifolia Salisb.), §, phylicifolia L., E, viminalis L.

and §, caprea L.

A considerable amount of cytological investigation in this

genus has been carried on by the English worker Wilkinson (1954,

1941). His 1954 work is a preliminary account of the cytological

method of distinguishing Salix alba var coerulea Koch (§. alba

var. 23133 G. F. W. Mey.), the cricket bat willow, from closely

related Species, varieties, and hybrids. The work was based on

studies of root tips.

It was found that all §, 212E L. varieties and the hybrids

involvinglfi. fragilis L. showed a somatic complement of 76

chromosomes (i.e., tetraploid) and that it was therefore im-

possible to segregate the cricket bat willow on the basis of

;75-

chromosome number alone. Evidence, however, derived from the

examination of a large number of somatic metaphase plates of

the £123 varieties suggested that their differences might to

some extent be correlated with the number of satellited chromo-

somes.

The cricket bat willow showed only two chromosomes wifii

satellites, while the other 2122 complements examined showed

at least four. Wilkinson further suggested that there are

some indications that the nature of the cytOplasmic background

might be helpful in verifying the results arrived at on the

grounds of morphology alone. He found in this group, due to

the presence of tannin and like substances, the protoplasmic

background, with any combination of fixing and staining, is

troublesome. Langlet's modification of Navashin's fixative

gave best results. Heidenhain's iron-alum-haematoxylin was

used for staining. Drastic variations in mordanting and stain—

ing periods were needed to avoid over-staining of the background.

Further work on the cytology of the cricket bat willow,

which further verified his original work, was reported by

Wilkinson (1941). Individuals of the cricket bat willow from

a wide geographical range were studied. The satellited chromo-

some hypothesis was supported in all cases but Pratt's (1954)

suggestion that strains of this willow exist was correlated with

the existence of types with abnormal somatic chromosome comple-

ments. Wilkinson concluded that the cricket bat willow was a

variety and not a hybrid as described by some authors (see foot-

note p.62).

"74.;-

LITERATURE CITED

(Salix)

Barnes, H. F. (1951) Further results of an investigation

into the resistance of basket willows to button gall for-

mation. Bull. Applied Biol. 18: 75-82.

Blackburn, K. B. and J. W. H. Harrison. (1922) The meiotic

phase in the Salicaceae. Rept. Brit. Ass. for the Adv.

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.................... (1950-a) A list of chromosome numbers in

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--.................. (1950-b) A list of chromosome numbers in

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nigra and Salix amygdaloides tOgether with the hybrids arising

from them as these Species exhibit themselves in the vicinity

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-------------------- (1895) Relations of Salix missouriensis,

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157-150.

-------------------- (1896) Salix cordata x sericea. Bot.

Gaz. 22: 592-400.

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oploid, bisexual species. Dansk. Bot. Arkiv. 5: 1-17.

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viminalis x caprea von Heribert Nilsson. Hereditas 15: 1-52.

-------------------- (1955) Die Konjugation der Chromosomen bei

einigen Salix-Bastarden. Hereditas 18: 199-214.

Harlow, W. M. and E. S. Harrar (1941) Textbook of dendrology.

2nd Ed. XV + 542 pp., 254 figs. McGraw-Hill Book Co., Inc.,

New York & London.

-75-

Harrison, J. W. H. (1922) Interspecific sterility. Nature

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.................... (1924) Sex in the Salicaceae and its

modification by eriOphyid mites and other influences. Brit.

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................... (1926) Heterochromosomes and polyploidy.

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(Tokyo) 50: 516-520.

-------------------- (1918) On hybridization of some Species

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entry $680. 19 )

-------------------- (1922) On hybridization of some species

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artificial interspecific hybrids in North American forest-

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angiosperms. A mer. Journ. Bot. 17: 152-174.

Lotsy, J. P. (1920) Heribert Nilsson's Onderzoikingen over

Soortsvorming Bij Salix met 0pmerkingen Mijnerzijds omtrent

de Daarin en in Publicaties van anderen Uitgeoefende Kritiek

aan mijn Soorts-definitie. Genetica (The Hague) 2: (162)-

188. (In Dutch. English review pp. 187-188.)

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chromosome numerals of Species of British flowering plants.

New Phytol. 58: 1-51.

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cious plants and their relatives with special reference to

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(Cited from Gaiser, 1926.)

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Nilsson, N. Heribert (1918) EXperimentelle Studien fiber

Variabilitfit, Spaltung, Artbildung und Evolution in der Gat-

tung Salix. (Experimental studies in variability, Splitting,

Species formation, and evolution in the genus Salix.) Lunds.

Univ. Arsskr. 14: 145 p. (Bot. Abstr. 12: 670. 1925. See

also abstract in Genetica 1: 555-557. 1919. Reviewed by

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-------------------- (1951) Uber das Entstehen eines ganz

Cinerea-ahnlichen Typus aus dem Bastard Salix viminalis x

caprea. Hereditas 15 (5): 509-519.

-------------------- (1955) Die Analyse der synthetisch

hergestellten Salix laurina. Hereditas 20: 559-55. (Plant

Breed. Abstr. Vol. 5: entry #1055.)

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Salix. Japan. Jour. Bot. 2: 1-55, illus.

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. ry 28: 12-20.

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2nd Ed., XXX 4 996 pp. MacMillan Co., New York.

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XIV e 574 pp., 48 figs. Princeton University Press, Princeton.

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in Morus alba and Salix amygdaloides. Ohio Jour. Sci. 19:

409—416.

-------------------- (1955) Observations and experiments on sex

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8: 202.

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A. Yearbook 1957: 1241-79. 14 figs.

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-------------------- (1929)* Chromosome studies in some

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forestry. xxxvi + 455 pp., 140 figs. John Wiley & Sons, Inc.,

New York.

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Kreuzung bei Weiden (Salix). Zfichter 1: 125-126, illus.

-------------------- (1951) Zur Verbesserung der Schalweide

Salix americana Hort. Der Zfichter 5: 569-570. (Plant Breed.

Abstr. 2: 140. 1952.)

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-------------------- (1941) The cytology of the Cricket Bat

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of the seeds of aSpen and of some species of willow. (Russian)

Proc. Imperial Forest. Inst. 11: 269-274. (Cited from Smith

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vy, J. B. (1955) hale trees of Salix alba var. coerulea.

FOLCStI‘J (3: (IS-EV.

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Soc. Botanioue 1‘ru1aoe c9: WS-olc. I I

--- ------ --- (1925) Le characiere tire de la Eragilite dos

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75.

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out. Wendrol. Gesoll. 58: 57-59.

x}

9 1

GBr" R ( 92o) Salix copusiensisOlosz cz'M und lore Eltern,

b

.

S. Litalbeliaha ‘..illd. und S. (31311110119. L. leagyar Bot.

LapOk 25: 95-201. _ ‘—

-------- (1928) Salix silosis ca "illd. und ihre dyer

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---------------——--- (1928) .911x laurina. Die Entwicklung

und die LBsung einer mehr als :1undert .Hhrien Phlegonet-

ischen Streitfrage. Lunés Univ. Arsskr. (n. f. ) avd. 2, Ed.

24, E0. 6: 99pp.

-- ----- - -------- ---- (1950) Salix cinereax Utbredning och

Ekologiska Detingels er 1 Sydestra SVcrle. Botaniska Eot-

iser 1950:129-145

------- ------------- (19C50) Dis trinution and 900103y of S.

cinere in sodt1ern SNe den. Bot. Eotiser 1950 : 129-145.

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some DdllXo (in Japanese) Pot. La ". Tokyo 27: 185-195.

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alsiya Ann. Soc. 2001. Hot. Fenn Vanamo 5: 155-275.

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101: 851-861.

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low. Jour. of 15 ot. 74: 250-55.

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5_‘alix. Leitschrift Induktive Abstsmmungs u. Verbungslehre

5/. O49_OR

qr

'thm

'(

CC) 0

Sugiura, T. (1:53) A list of chromosome numbers in angio-

Spermous plants. 11. Pros. Imp. Acad. Japan 12: 144- 46.

- ------ ~--- (1958) A list of chro~osome numbers in 93:10-

Spermous plants. V, Proc. Imp. Acad. Jgpqn 14 (10); 591_

592.

---------- - (1940) A list of olromosone nlmirs in 21io-

spermous plants. Proc. Imp. Acad. Japan 16:15-1g

Sukaohev, V. 1;. (1959) Selection Glipel 17116911119 ith Willow.

(Russian) snoe Ehoziaistvo 5: 24-54.

Vamazaki, R. (191') Chromosome numbers in some angiosperms.

J‘J‘i‘. JOUJ’. \ICI’lCTt. 12 (5:5): 101-1037.

Yampolsky, Cecil 4 Helene (1922 ) Distributions of sex formsv

in the biomeHO‘iii flora. Pibliotheca Senotica 5: 1-52.

3779115311714"; C KAI-3 Lind 1 e ;r

Kalnut bamily

The walnut fawily includes about 40 saecies in 6 generr

distributed thruout the temperate regions of the northern

hemisphere and also, limitedly, in certain tropical montane

regions of botl hemispheres.

Of the 6 genera (Jua‘laus l1. (talrut); Carva lfuttall

...,

(hickory); Platjcarya Siebold and zuccakini; Pterocarga Kunth;

Engelbar tia, and Alfaroa) only the first two are native in

Eortb America and it is upon the valuable and useful timbers

and nuts of these two genera that the importance of the walnut,

family depends (Rehder, l94o; Pearson & Brown, 1952).

JU9LAES L. (walnut)

The walnut genus includes aoout 15 species of trees1

found in the forests of North and South America, the West In-

dies, southeastern EurOpe, and southern and eastern Asia. Of

t-i

. cincr (Dthese, 6 species (J. niara L., a

_+ I’M

., J. hindsii (Jep-

son) Jeoson J. californica S. Ratson J. runestris En elmann1 :_ :_ 1 L2 )

and J. major (Torrey) Heller) are native of the United States

but only J. rivra (black wa‘nut) and J. cinerea (butternut)———-M———_ —————-—-———

are important as producers of lumber.

Most important of the exotic members of the genus is the

cosmOpolite Juglans rewia L. (En lish or Persian Walnut) whichL4

Ho

8 the source of the French, furnish, Italian, Circassian ano

Indian walnut timbers.

l Rarely shrubby.

J

ialnut, esyecially black and English, is the lea i

C5"!

P

‘

wooi from hfliCfl gunstocks are made. The timbers are also

2).)

prized as fine cabinet an furniture woods.

The nuts from the various species of ualgns are

rich in oil and nutritious. Cf special importance are

the English walnuts of g. reria, black Walnuts from g.

nisra and butternuts from g. cinerea. It has been the

direction of man's interest to the edible part of the tree

Which has resulted in the considerable breeding work car-

ried on in the genus. (Harlow & Harrar, 1941; Brown &

Panshin, 19403.

ARIIPICTAP BafiaflTitn PRCH nILn 8T03K

With the possible exception of twe genus Carya, no

other strictly forest tree genus native of the United

States has been as carefully combed for superior wild forms

or varieties as the genus Juglans. The stimulus for this

selection effort has not, however, arisen from a desire to

isolate superior forms for forestry purposes but rather to

locate better nut producers for the more immediately prof-

itable nut growing industry.

For the most part, selection has been confined to J;

nipra and j. gingrga--fortunately the best nut and timber

producers native of the United States.

Early interest in the selection effort is traceable

to the "nut contests" sponsored for many years by the

Northern Nut Growers' Association% aided at times by the

1

*Established 1910.

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enus b~r root ~r sues cuttiz 3, treated u n auxin or un-

(V)

treated, have not sire to the aitaor's attention. This neg-

ative evidence thus supgorts tee general coneeution that

past root’ng methods at Least have been unsuc'es ful.

Investigation of rototing possibilities

cuttings (f the malnuts wouli a;pear to Le a most valuable

at 11']F}!

line of investieati n. For forestry purposes the Lul

(n

or grafting techniaues are, of course, orozibitively expen-

sive. Jith a rerfected method of insuring hih rooting per-

centage of cuttings many valuable Walnuts now in existence

could be immedia tely utf.lized for forestry pur;08's. I

particularly promising forestry breejing possibilities in

the genus til , for the present, probably be neglected in

favor of those cehera thich may be cheaply propagated asex-

uallg. It is to be hope; ciat more long-raige government-

financed forest tree breeding

tainéd at the Institute of porest genetics, may be fouajed

for work on such genera. The breeding and selection required

for pure line develrpment DI superior stock so that reproduct-

ion by seed is fossiole is avowedly a long-time project but

is certainly Worthy ot practical consideration.

A possible short-cut method for the synt-iesis of true

breeding forms capable of consistent reproduction by seedage

in this genus apoe 1rs gnwmnbsmyt‘uith perfection 3f the col-

chicine technique. The creation of auto- and aliopolyploid

forms may ”Ti ld totally dif ere nt aii superior varieties of

[.43

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CW

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,- - .LL,

.5. ..L one

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f“

y- . ' ‘1 P‘ i - .~. 1:) l m ‘1 1 e h .3 I \ - ,7 h l .‘N I r .1, .- ‘. v ‘. 4»

t - J.- - l M L 1:) V .A. '— L-i {.x w Cd L) .. 1;; VJ .. » .a L.) L- ..z

N” a H, a , -‘..‘JbruiQSOLWITfUKCSr

"r“- .- -.. , .9 .-. ,4. V- ,1. , .1. , .. ,.M.'-4. .1. ' '1, ....-. - ,, -' 2'1 ' 3 ..j

lfl‘ calllufie CIIUWUBUME CUJUU 13 du.laub lo ”nut YLCOIQ-

ed bv aaococr (1:15). This worker reportei tne unreduced num-

ber of J. californica S. “atson and J. c. cuercina babcoca to

be 34 which would imcly a haploid number of 17 chromosomes.

A study by aoodworth (1930), however, of meiosis in 5

_ ' .., . 'r 1-, ' — ‘ ,1 , H

. TJQCSlTla -Cictu., o. n_ r“ 1., J. Cinerea L., J. t~.1°hur:ca

Laxim., J. sieooliiara var. cor iformis (iaxim.) Lahino, 31d

J. not a Feni. (— J. sieboldiana X reeia)) revealed a reduced

number of 16 chromosomes. meteroaloidi has not been reported

(T)

('9'

('1)

r+

H 53.3

rL-‘

H O H.

U)

m W a >I

CJ

("I

C H F].

O m L.

U

k:for Jualans although tin

noodtorth (193?) in Sarva.

Cytological stuiieg

goodworth (1330) reported meiosis normal and regular in

1 hybrid listed above. goragoloyigalry ug—LHE 806016‘ 833

(D

,1}-

L4-

D D CI

R:

b I"

C‘L

m fi'

1...:

117

p U)

(“I

SO

r—l

i C *fi

iform microspores were arojuc

these soecies.

.0 - 1.. .,_',I ~,:.,:., -7».- ,.— . . ..., .2

J. notna 0; thumb nvsiii Ulafljfl ‘nouei arEut abnoraality

u

v

in KiCYOSQOTOE€?€‘J‘- Althougj pairing at diaainesis appeared

normal, metaphase I shoted great confusion. baby bivalents

separate; and passedAtneir poles without Iirst moving to the

1

mid-1e region of the spindle. retauhase I Was t US, in efiect,

”W

eliminated. Division II was also marked by irre:ularity.

.,

Fl {7P!

.A t_

#4 v

’x

‘v

31)

O t_J

.

n Y O "(3 ~.-.’— ,

" at“. i '3‘ 'ir‘\

IWTve_TfiE PI”TD

(AT _~lons)

Au_stin, L. (1933) Science sceas new trees for the foncsts

of the future. 24pp., illis. rltcerville, Calif.

Rabcock, E. P. (1910) 'Malnut-01k hybrid experiment.. Amer.

Freeders n39. l (5): QCTO-B 2.

------------- (1915) Jlflles in Juglans I. Stacy of a

new form of Ju9lnns califor ica Tatson. Calif. Univ.

Pubs. Agri. bci. 2: l~«£.nl915.

------- ----- - (1151 4) {Studies in 31191-9113 II . Furt‘r;or ob—

SGI‘VEthl’lS on 8. T1611 V:-11’ 1.31;"; of T11.T_:.ns (3: Tji form? C 1,3931901’].

and on certain supposed uwalnut ork "merits. 38111. UDlV-

Pubs. flgr. Sci. 2: 47-70.

---------—--- (1915-2) A new walnut mutant somewhat sim-

ilar to live oak apIPr 9 in California In four different

localities. Jour. credity 6: 4L-450

-—-—--------- (l9l5-b) Uslnut mutant investi99tions. Eatl.

Acad. Sci. Proc. 1: 535-537. .

-——-------—-- (1916) indics i Juglens I. (1) Further. II

evidence tht tie o'Lll‘e waln t oriaf‘ates by putation;

ii

(2) a f>erollel emu!tion in Ju9lsis bin 3 (Jepson) Sarg.

Calif. Univ. Pubs. A9r. Sci. P: Vl-QU.

Brown, it. P. & Pans?d11,fb.l. (1940) (Run Ierc101.iorr'ers of

the United States. 1st ed., X3I+-554 pp., 512 figs. RcGraw-

Hill Poo‘c COUppr, Inc., hew Yonh and 10ndo

Crane, H. L. (1937) Nut Breeding. U.S.D.A. Yearbook of

Agricilture, pp. E~V-L‘90.

est, E. b. (1940) The distribution of self-sterility in

the flowering Plants. Proc. Amer. Phil. Soc. 52: 449-518.

Harlow, T. M. and Horror, E. F. (1941) ”TexWibook of dendrol-

05y. 9nd Ed...V+ 5”? pp., 534 1193. icGraw-Uill Pook Co.,

Inc., Lew Yerk & Londor.

Johnson, L. P. V. (1959) A Gas

artificial interspecific byh

m (o

riitive list of rttaral and

c in Northimerican forest

tree genera. Cancd. Jour. Re . l7: 4ll-~:44.

llotzsch, J. F. (l554 ) Ueber due Rutzantcndung der Pfanzen-

Bastards und his cilinge. Fer. Verload . K. Preuss. A{Qd.

Wise. Fcnlin lQSA; .“S-So

Pearson, R. S. 9 Brown, M. P. (1952) Commercial timbers of

India. ”alcu,t9 Government of India Central Publication

Franc“.

Reed, C. A. (1926) Nut-tree ocooa9ation. U.S.D.A. warmers'

Rul. Ho. 1501, Lash. T. 9..L/ J’

Rehder, A. (1940) hanual of cultivated tr es and shrubs.. _ f '-' u ». v C W ,9 . ... y

2nd Ed., 9~o pm. FQCullifin Co. low Yorl.

rees of torth “Movie§

(’1

CCD

—f-

b

L-

o

q

-4b"

o A

H

.(‘J

N)

v

5‘

..2

3 }.J

O ”'3

Ct

'0

Schus er 3. h. (19'?i) Lifferenccs in heterosis of walnut

" o r)].b {317.

010 C. (1941) boocies hybrid

" IQ -- , A -

SWitt. C. L. & OM8P3, L. Q. (1)0) b“"tlor stuCies on

pariods of recoutilityeand pollen vicbilit y in tbe pLC'L.

Proc. 17th finn. loot. fox. Pecan W1 £83.: 77—95.

(“x

'T‘Wgu'), F. p. 7k “OWT'WT!” 1,. 7:. ( Aft?) I; JET CWT" Clip 01 lTT‘Ul 113’;

rollinwiiov in tbe 1ecwr J. £9"ic. ?1s. ’". 9‘7-9;.

Toad . T. (1“‘TT NTJD’wi“. «0111H01ion es s Fee's of in«

-‘r:sin| neon tion i~ CO"”€H019T vvrrien 1'lnnt o'chfirds.

”0c. “'gfl. foo Port. Sci. 99: 104-66.

-----*-"'—- (1935} POTllMHtTOK ~10 Llooni= }”Lits of t“e

“cesiwn malnut is 3‘1f9r11i9. Cock. full. U. 3. Left. A9~.

To. TNT: ff. 53.

‘”cllfi-:9‘mn* E". (11531) ureevi1n_;stl111ts. P1T13. 919'. VIN; 3”.

Ass. 99; 15—21.

Toedro;t3, T. H. (17"9) Iciosic of mi910990309enesis in tUe

Ji‘ltr‘_-cf c. 1¥;r. Jon”. 9st. 17: 563-569.

‘f.bIo;OV, R. L. (3’76) Inte?—s;eciws Tyoridixet1or of walnuts.

Soviet 51 tr19ics KO. 5 (71): F5~99. (Cited fnhw Jobnson,

lotio.)

LIL“ LUJ”TT T133 GILL“)

(.N"l:rs)Mb-..-—

Anonymous (19lo) Lutet1ors in walnuts. Jonr. credit; 7:

1:517U. .

--~---—-- (1995) ”HlflUt-U‘i9‘ . Introdwction of iwm1ne ver—

ietj b‘ tie department. “CH 4691. Jonr. A93. 97: 25.

3.1731313) LlO"LT (1:127) [31. UCW’ erltET‘nPi Se in joce St tree breed—

in;. JCTIF. Eanssfrj'295: 92L»5-;Lm;.

--—--———--——- (1928) Vtecding pines for mone repii growth.

Jcn1r. “’ciwfitii'y it? ('7): Rani—3K1}.

-—--——---~-—- (1998‘ Bx1e”1tents it the Eddy tree breeding

stcticr. Ti‘Mergwr 99 (7): /9 34

—--—————~--—- (1999) Ike Eddy 'ree Prcgdin, Station. 19d-

rono 1 (l5): 05-.7".

--—-~—-~——-—- (1932) Pine CNO vzlnnt wrecri1* for timber

p?oJUo9502. PWOD. 6th Intesztl. COD . Genetics, Itfz;9,

1952, 2: 2-9.

---———-—-——-- (1977) TFO institute o? forest 9cnetich. finer

IForests, September, 1957, pp. 5.

~—--————---—- (1958) Forest ‘ oeti1.. .u1loson1D Report

1938: 435-49.

Peacocw, E. 9. ( 910) LT:tOLO09y in Jn9lnns cslifornics ”et-

son. Plart World 13: z.~31.

Vstctclor, L. D. ( 915) Tsriniio: 9rd Ili9tt resistincc

8111011.", :.'.'="1:',_,111'.9. Cilii‘. 13,".th 301.121.. Fort. "011t3113’ 711.11. 4'

4“b-450.

-—-——--—--—-—-—- (1916) B“ohlens in walnut orceding...usl—

nut blight and var19b111tr of the present Iroves. Jour.

1931edii“v 7: Gi-LIE.

Efisse , P. (1914) The James river 1191:1111.Priormlly crcss

igetwween TfllttCIThlt T‘UT 5‘291191‘1J.ljn1t. Jour. EiergM-t7 .g

01(1

Biibv 1. a. (1919)

Amer. Nut J01c.71W

.3; .’

jEurbank, thEcn. (19 o) Tzc tixdni

Trail}: royal 3.1""! {7 , a 1- ' TJZTJV‘TTC 1“

profit...8 $9., 111v.. iant. H

' e, 11t't<3:11111t

VU-Hr C

3

HBO"

'7

La

1190 {‘01) I

11f“,1

"TJ

‘ 7

IT 5: V. 1

1nut.

.1..

is r;; the

.L.‘ t. . __, ,. -

Lntbca1t; 9nd

P11! .1"...

P1J1Tw1nli, ILJtEuJF (lQlr ) f1<: LW1Il1erfl: *«119ZL 1 lYPlt I3icn1€<3r

jest. LHWUevmnn "l (P) : Pl.

CQfiTCT‘, IT. fl. (JLXMJ) Studies ijlivxzediay'1.,3ffib1strotefl 1;?

1iie trixfi<:xmes of‘:figecies 1111 vv1rdhls of JW;fl’”1°, an11tFCP3,

P°1oveW, 1nd (olsvvm o7 pp., 11143. .'sblgguon, L. 3.

"o1“?1‘f, l. 1.. Tlhl“) ~n1 flhefln°nt'.w,lTHL. llrr“ Visit 1%:

1’ ‘k‘f;

u“ 1..

I11vis, lfl. F. (liHNJ) 19x; toxfix: priruziplfa of JWM lots r1_‘vs

qs ideni;1jied with synthetic JHQlone, so" its toxic 3.lects

011 t‘on11:0 :1n <1ljkiliPI 311-11-11. 2u194r. J“11:‘. F701;. 153: Sill.

He: s.oe~5 J. I. 11757) StCWtfinv 1 jPClQPCQd str11¢ of tiWPcr

and c:mms tre es ir11fiw1 fOrestry"“wafirqm 0; tle WP” ”(s'ec

‘fs'llxajr ‘11‘11C1 i-'t31. 1111 11. C<Tr”v:nlf;. 1(31111'611 1511; 711'. 1-s:3. ‘ e11t

-P--15, ll pf

Frr~ki;;s, 11. 13. F- Smuiwcnfith, 1}. 1?. (17197) anno; b 01"the: cont-

1 ’ ’1h1 not 9nd otier e '

. ()Ofl...

O

». _ n '1’

,-" _ Q .. - . 6.4 A-.

Jepson, N. L. (1909) SPonteneous hybrifis Of native Cfillf-

01111911'trecw3 flung“. Eqn336<1°s' 1J39001 }?e;fi.. E: Rift-cl.

h1rdisti, 8. ( 93 (L lection of 1cluuts) Soviet Sub—

trOoics N ‘ 3

quek Hslnwts of Iowa. trans-

Ions St. Nov,. 5 0. 7c: 2 O-l23.

Rebel, J. R. (1911) CyiolongOl espoots of tro1lin3 nuts.

Broo. It?» Phlt ?v. $3.. 5&3: 22—24.

FEIFGP E. .. (1937) Toces on North American trees ang

.1» 9; '- '8 A _ VIII

814l'11133. J. .:-.111. 11111.). .LOZ 100-1110.

Pr,..'."1’3"1 ' 1 1.1. wHo h 1:1' 'V 3 o -1- o (191(3) A. Vq I i3 t _, 1n tne ulsc{—1.

O :3

walnut. Iowa Aefid. Sci. Proc. 25: 241-42.

----——--————---—- ————— ———-- (l9PC‘) A varistion in the blao

C;

walnut. Proc. Iona Rood Sci. Pfll-fe.

PlllCUOt, C}. (ngDQ) E?opo;%: of tire ccnmWitt 13 on Lawsodjjg;

forts t and nut trees. Amwr. Preeders' Ass. P01 3: 304

Ell.

Reed, 3. A. (1955) Turko"s, men and nuts. Jouy. Nered.

O

t o

0

7L- ".

2[L: L)L_)tJ-E'().

(— I7 ‘5 . '1. ..J. 7.. -l- - ’ " - ,.

-__———---—- (lflop) LQtQPQl Mflluho LZUPIJS in the Ofist.I

F) ,. 3.. ()r -‘1‘ 7" -1. 1171.11, 117... - #1-, A ‘ f ,

ref). l-'OCo “it. “Inc 11121. 1': LL. 1.1.14 -171). 1318300., me_1_3‘.'6,

159;) A tybrid walnut tree. Gard. & For.

Fuldhcnét , '3. 1 . (1&(Y7 I‘e11C1't CH” t313 Conunittxse CH1 bITN dirk;

ANT and OtHPP forest trees. Amer. Byeoders' ASSOC. Rept.

3: 224-25o.

-“"-'“’““-“-" (1909) Preliminary report of the Chairman

of the committee n treedin; not and forcet tree . Amer.

F)

Breeders' Assoc. Rept. 5: PBF-PS-

---------------- (1910) Report Of cowwitteo on breeding nut

and forest trees. arer. Freoders' L93. 1 (3): 105—193.

-——--—-------~~- (1911) (Same tith as above). moor Ryo;c-

ers' Ass. Peyt. 7: 250 959.

---——~—------——- (1919) (Sine title as above). Amer. Freed-

ers' Ass. Pe “t. a: 515-522.

VOlZ’ 9‘ 3' (1920) The Edd? tPOC breedins station. Calif.

Countrymsn 14 (6); 8-15.

,

K

k

Carve (= fiicoria Paiinesoue-Schmaltz) is a typicallv

American genus since 20 of ‘he 22 species are native in east-

ern North America, south to hexico. The other two are native

to China and Indo-Cbina respectively. The genus is divided

into two sections: (1) Aoocarva de Candolle (pecan bickeries)

and (2) Eucarva de Candolle (true tickories). Roth sections

contain a considerable number of Varieties and hybrids which

are sometimes difficult to identify.

The genus Carve is the source of the tough hickory Wood

of commerce which receives world-wide use as handles of "im-

pact" tools sucb_as hammers, axes, and picks. In addition,

the wood is extensively used for various sporting goods (skiis,

golf club handles, etc.), furniture, ladders and numerous

special products requiring strong, tough, elastic wood.

Most important of the commercial nut producers is g.

Eecan (Marshall) Engler & Graebner and its numerous named va-

rieties widely planted in nut groves thruout the southern

states.

As in the genus Juglans, the interested breeders in this

genus have been horticulturists intent upon isolating superior

nut—producers rather than forest tree improvement.

ARTIFICIAL SELECTION PR m WILD STOCK

Parallelirg the work in Jumlans, considerable selection

effort has been directed to the genus Carva by nut growers of

the United States. Interest for the most part has centered

’1 a / ,-_ ,. :" ‘ O ,. " °

(oarya p-311 (Lire-a11) Lflilcf‘). cele3t1~' for n:t 1uuiity

' . ‘\ T‘ . *- r \ .. v ‘ 'u ‘1‘ 7‘ r

nae als- 1eun 3afrl- on hicJ ouciess 1 on: natural loims

(Licnaux) Louf. (shellter: hickory), Q. tomentosu (nam.) Kutt.

kD

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selections for nut gualitt up to tee time.

Letters of tne genus Carve a) are tly nyb-iiize with

ease in nat11e (orgne, .~37; Bender, 1343). Jennson (1)39)

listex 7 natural hroriis. The cpief source of 1nd.“Dati—

robesly traceable to variations

1n flowering time there tne ranges overlap.

The 11 teriture revels no account of artificial selec

tion fro1 Wild stocks designe; to isolate superior forms for

strictly forestry purposes.

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A la year ulz JEJ Q variety stu 1;; ex HbvhrOOl

r3} '. — -. 1".“ -' M _ - K . ; I'm: m-L .~ ‘ ‘ 4-.-

541 ELlUutS vu;iCL.(:Olc L¥r,3;5 cxzeains. 'Fue txlua

L)| ‘ " " ‘I '~ ~ " I' ‘ r 0" ’ ‘ .‘ u’“ “. " . A‘

_aac; antler Cufltkjfifl an tie aveiags 8,Lu4 dull

JUECET 01 pallen gra US proauce;

(”\l‘

tree tnus arounte; t3 aggroxiuately a;,394,llj,

u. v‘ rwl\I (’4 n . I‘

.. P

- ‘ — ‘ A F“in; at lC(3 efTiCfiTHCV: 106': lJ eacn H’LLCH Cr

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t at tale collen MUJL- uaVe be m Sdl‘lCl‘l ad

%? ie‘ Ev Var101s “OrLeTS- 311 e (1957) In flea

tgat iability of fecan tollen varies greatly u

4‘ .

seasonal confitiajs a1: otler lactors. Neodroc

J

stored polleh of 20 varieties unfier various te; ..4

l

h‘midit coafiitions but osservei that 'no temps

kuxiiiity conditiLH1. LS founfi under VfiiiCJ pollen

storei for longer than 96 hours (4 days) wits c

germination." Smith & Bomberg (1932), however,

-men of she ;ecan varie

O N (O

r’.

C)

H;

(<1

0 C)

H.

(+-

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pollen of another variety remaimeu viable for :

,+- ‘4,» -m,.-.:..',.—,. . 41,, .\ .-,«."; -

vablons MlLu wu er Valltelco um has chdfl anl f

tainei for 10 flavs or more.

(l 30) had

l number of

ain were in-

J

r com3ute;

prOiuce 229,

has been invest-

itn varieties,

f (1328)

wii('0"' «‘4' ~

gerauur

rature and

ertainty of

resorted

t7 under lab-

11y 8 days.

tfieir Cheer-

canal that

ll be main-

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17301153“, I}. P. C: P91103111, .‘2. Jo (1.1.1-1 1 CO? ”CPO-19$ 1J1Chg Op

_ o -3 1 L ‘4 '7‘t(\ 10, .. ..

he I11ted States. lst eo.,KI+-EF1Q pp., dis iigs. Lo3r2d-

T v r T.‘ r 1 ' I A. ‘ru .‘ fl . . "t . -.1

11111U 1001 Cenfi1-n3, Inc., Lea ,Orfi awn LOnuOn.

v '~~— ‘ 7-. 1 *1 1 xx... 1 :1Crane, n. L. (lJo ) Lot bPCBLlLv. .S.D.m. ;earbooz oi

1.1r1calture, pp. oev-ee .

Fork1rt, C. (1914) Taelve years' experience in l1ybridizing

pecans. Natl. hat -}rouels' As soc. Proc. 13: .t-"O

Harlow, T. 1. and Larrar, E. S. (1341) Textnook of dendrol-

OfV. 2nd Ed. XV+—542 1p., 934 figs. chraw—ill Rook Co.,

Iris. , 1 e17 ‘KOIQ: f: llorui(12.

Pearson ET. 8. ,r_. Prown, H. P. (1932:) Cormorcia l tin1.1ers of

Indix.. Zolc1lixz,.rc1o1wmwx1t of lrxlia Cerfluflil l‘uol iCXFtiOU

Franco.

Herder, A. (1943) Lemuel of otltivotoo trees and strvbs.

2nd Ed., XXX+-996 pp., FacLillan Co., Fe York.

Pisien, E. 9. (1914) Tree breeding. Natl. hut GPOLOPS'

Ass. Proc . 13: 50-33.

Serg:er1t, C. S. (1922) Annual of trees of Eorth Arorica.

Sno Ed.,1.ew York.

Smith, C. L. 7: Pomberg, L. D. (19"?) P0llination control,

peTiOd oi receptivity and pollen vietility in the pecan.

Yatl. Pecan Assoc. Bull. 31: bo-Ul. ,

-----------*-----~----------- (19?3) Furthnr studies on

periods of roce1>tivity and pollen viability in tne pecan.

Proc. 15th Ann. 1 eet. 3ex. Pecan Gr. 38s.: Vo-e

---——-———--——----------—--—-- (lV40) Stigma iecept jtv

and pollen sheddinq in some pecan arieties. J.1Agric.

680 CO: 13‘7“"‘Vro

mnqhfi, {1. P. &;Emynmcr§, l“ 1). (-9">7) Duetbods Of'anrtrolling

pollination in the pecan. J. Agric. Fee. 47: 267—

Woodroof, H. C. & Pailey, J. E. (1928) Unfruitfulness of

the pecan. Ga. Exot. Eta. Bull. 143, 40 hp.

fioodroff, J. G. (lQEO) studies of the strul1ate inflores-

' genes and pollen of Eicoris :ecsn. Jour. Agr. Research

40: 1059-1104.

Woodworth, R. U. (1950) Meios318 Of miciro.po'

Jolendacex1e. 1mer. Jour. wot. 17: 865-8’

‘1ogenesis in the

C)

C)’

Lll‘RAEUVE NOT CITED

(0 m)“&

Stucke;¢, H. P. (1916) The two groups of varieties of the

Picoria fecan and their relation of self-3+;er111t7 Ga.

Agr. Expt. btfl. Bull. 124, 22 pp.

Woodroof, K. C. (lQBb) Develooment of the embryo sec and

young embryo of ricoria recan. Amer. Jour. Rot. 15: 416-

421. ‘—‘

1 For further references not cited for tlis 'enus consult the

literature lists at end of Jurlans ane ctio' __.aé_—_—

BETULACEAE Agardh

Birch Family

The birch family includes more than 100 species in 6 gen-

era: Betula L. (birch), Alnus B. Ebr1art (alder), Carpinus L.

(hornbeam), Ostrya Scepoli (bOp-hornbeam), Ostryopsis Decaisne,

and Coerus L. (hazel). For the most part the family is re-

stricted to the northern hemisphere. A few species of Alnus,

however, extend southward through the mountains of Kexico and

Central America to the Andes mountains of Peru (fiinkler, 1904;

Sargent, 1922; Fender, 1940; Harlow & Harrar, 1941).

tembers of the family are trees or shrubs with alternate

deciduous leaves and monoecious or rarely dioecious flowers.

Except for the members of the genus Petula, little breed-

ing or cytological investigation has been carried on among

the arborescent forms of tie other genera. Considerable

breeding and selection, however, has been carried on in the

non-arborescent genus Corylusl, designed to improve quality

of the filbert nut of commerce. Breeding work of this nature

does not fall within the scepe of this paper.

In tre attacred "literature not cited (Retule)" will be

found references to the cyt0105ical researches of Woodworth

(lQBQ-a, lQBQ-b, 1930) in the genus Alnus.

l Corglus avellana L. of EurOpe produces the filbert nut but

the United S'ates' supply is furnished largely by locally

selected forms, varieties and 3ybrids of this species.

-1C.'-O-

I l L—4

r":

J [—4

o

A

O 1.1.

W O 3‘

V

The genus Betula is made up of about 40 shruboy or

arbore scent SJeCiES wi ely distributed tlruout the northern

He:1s211ere. About 15 soecies are native *0 {he cooler re-

0

gions of North America. Cf this number 10 are trees. about

5 of these ma; be considered of corxercial imgortance:

;. luten licnaux (yellow birch), é nn‘vrifera Larshall

(paper birch), B. gooulifolia Karshalt (gray birch), g.

nirra L. (river birch), ani g. lenta L. (cherry birch).

Lost important of the native ejecies is the comraon

nort. astern hardtood g. lutea nhich, it is estimated,

supplies about three fourths of all the timber sold as

birch in this country. birch is one of the most important

furniture woods of the Uni ed States ani is also used for

a great variety of proiucts, including veneer, railroad

ties, cabinets, boxes, woodenmare, novelties, ani other

miscellaneous products (Brown & Panshin, 1941).

Cf some commercial importance in murope ani Russia

are the "European white birches" (g. oeniula Eoth, é. gubescens‘———_

Ehrh. and their intermediate forms). g. alnoides Hamilt

5..

and g. cyliniros;achvs Garble are timber trees of com-

mercial significance in India (Pearson & groan, 1932).

Laiy of the bircnes are used for decorative purposes

in the United States. The handsome foliage ani bark of

s :ake them com on ornamental(I)

the EurOpean white birch

1 ' . 1Cther native birches of local importance incluoe: b.

gapyrifera occienfalis (Hook. ) oarg., var. western

oaoer birch); B. oa2v ifera keiaica (Evans) Henry, var.

(Kenai birch) of the Alas {an coast region.

-101...

trees of the eastern states.

The tenus Let la is notably polVWUrhis due to the ease

witn which natura interspecific crosses a;.arently occur. In

ition, oolyoloid sgecies, varietie , and hybriis ranging

from trioloid to hexaoloid have been recognized cyt01051cally.

LE little evidence of conscious effort

in tne United States directed to the selection OF superior

individuals fir forestry purposes from wild populations of

he native birches. ExcepMins to this, eneral conclusion

may possibly be found in those few instances there controlled

breeding projects have been instituted. Eresnmably selection

of the bes t tild forms available tor use as oarental stocL

1" s- 1' "' 1‘ ‘ " ‘V ‘N .% —.‘ A

has LBt; given Some corsioer at on.

)

tt :t;',n has been reporteulv direct-(I)

In Europe some so

(

1

ed to the qu stion of seeea origin (groveenanc 8 studies) in

an effort to isolate climatic strains of the important é.

verrucosa Ehrhart (2 Q. pendula Roth and g. gubescens Ehrh.

ani their intermediate forms.1) Schreiner (1957) reported

German seed origin studies with a. verrucosa underway at

the Forstbotanisches Institut , Technischen Hochschule,

Dresden but any results of this work have not as yet been

reported.

l

The taxonomy of these two Species has been consideradly

contused in the cast. The natural more northerly range of

B. oubescens Ehrh. overlaps, in many districts, the more

sout~erly ren:e of §. verrucosa Lhrn. (Jensen l957). In

these area interteiir e firms showing a gradual transi-

tion between the two

0

sm»e ies led Linnaeus ani other early

taxonomists to gr the o suecies, JlUS the intermedi-

ate forms, into the one ecie s: g. alba L. (Congd p.lO53.

nettstein and Prooach I939) of he Kaiser nilwhelm-

Institut fur Luchtunisforscnung, drain baur-Institut,

lunchberg in an essentially cytotaxonomic report on g.

pubescens and g. verrucoca reported plans for the study of the

economic value of the natulal intr1e11te forms of these

species. R sults of this study have not been published.

Heiberg (1937-1939), in a report surveying the provenance

problem involving the important forest trees of Norway, em-

phasized the iaportence 0: select ng individual birch trees‘HJ

of good quality for further breeding, rather than attempting

to select clitatic r;ces Considering the natural occurenceLa

of large hybrid pregenies betwee1 . oendula anl é. augscensIt“)

such a suggestion appears sell-founded.

Sprangers (1937), wit; reference to the selection work

b lar invest gationsI...)

H O :‘XJ

(T

U)

H :3

(D)

‘1

{:3

:3

LA

EN

*1

(I)

O O

3‘

(I)

“1

(I)

}._lo

1.}

Fl-

1

to be conductea in Hollani.

In View of the ease with Which many of the species and

varieties of this genus hybridize in nature it may be pre-

sumed that there exists in the mild stete a wide variety of

segregates in F and later generations. It would seem ad-

2

visable, therefore, to conduct systematic surveys in regions

of range overla: of several species. Selections from such

wild stocks may yield valuable material for forestry pur-

poses which could be utilized immediately if vegetative

Ine work of Lorgentbaler (1915),

1:2 5), sinkle r & Anton (1933), and

Jentys-Szaferowa (l938)l1as proved the incorrectness of

this View ar 1 t*1e final se;yre 3 ation of the two species

and their in rnzediat.e forts. menier (I940) believed Q.

verrucosa Lhrh. shuld instead be csflled B. ‘endula Roth.

Footnote of p.|02con{'dd:

Helms and Jorgensen (1“

propagation tere feasible. In any case, the material might

grove useful for further breeding nork.

1'."I~hY-t\- ~7- 1. 5 :'1 ...,‘.,.‘.I(.‘ .“~ “ . .1' _,_""_‘.T ‘1’?

.JLVJLTKU 4.1.30 ..J_'£.a.v.1,1i'~.u 3.12 grudge/lieu

Although the incidence of natural interspecific hybrids

in this genus is quite highl, reports of breeding studies

beyond tde initial steps of provenance studies and individual

selection from wild stocks, as noted above, are reoresented

in the literature by only one preliminary hybridization re-

port.

Smith & Nichols (1941) reported nine crosses, the pro-

geny of which have been tentatively identified as hybrids.

The parents included: a. mandshurica (Reg.) Nakai var.

ienonica (Miq.) Fehd., a. kaximowicziana Reg., a. lenta L.,_A.K/

44" L

. niera L., g. populifolia Narsh., é. lutea kichx., E.

lm

apyrifera harsh. and its variety henaica (Evans) Henry.*6

Tests of vigor as compared With the parents have not as yet

been reported.

Flowering data

Schreiner (1938) reported that studies of the bloom-

ing habit in fistula was complicated by successive develop-

ment of flowers on individual aments and the aboarent abil-

ity of some species to mature parthenocarpic fruits. Smith

& Nichols (1941) reported species of Letula as extremely

protandrous, in some cases so much so that isolated trees

bore almost no fruit. There is, however, according to

these workers, no difficulty involved in getting selfed

(Johnson (1959) listed 13.

—104-

—. I- 0, Lb: :.- .\-.“ °. h :F. "7!: - ~~ : ‘ “

seed by artillCldl yblllflétluflo lfllS IS suuportei oi Last's

(1940) conclusions that the genus is self-fertile.

hybridizing technioue

The "greenhouse technique" as described by Yanchevsky

(1904) for Salix aid nettstein (1933) for figgulus, has been

successfully used in the betula crosses reported by Smith &

Nicnols (1941). These latter authors also noted that sausage

tubing may be more conveniently used in betula species(;n-

stead of paper or glassine bags) to cover a large numoer of

”‘1 ~

flowers on one branch. They recomtenle' plugging the end of

the tube hith cotton tool, thus insuring EOUd aeration.

(For details of the "greenhouse technique” see Salix section

of this report.)

Pollen longevity

Peoorts of pollen longevity studies in this genus have

not some to the author's attention.

It is probable that refrigerated storage of 2-3 degrees

C. and relative humidity of approximately 50% will Success-

fully preserve birch pollen for reasonable ser‘ods. In cases

where pollen is desired from remote sources it would seem ad—

visable to obtain cut twigs searing male catkins in the dor-

mant state and hold tnem in refrigeration. They may then be

forced as needed in the greenhouse. This method was resorted

l)

by Heimburger (1940) as successful witi speci s of Pooulus.

tenbers of the genus :etula are grouped among the

l_J

"difficult" o.ants to root from stem cuttings. In horti-h

-105-

cultural practice buiVing, g-aftins or layering is sat-

isfactorily used but such methods are obviously impractic-

able for forestry purposes.

The tort of Afansiev (1939) demonstrated that rooting

ability of stem cuttings in this genus may be significant-

ly increased by auxin treatment. Greenwood cuttings of g.

‘1 .C‘

populifolia Karshall treatet lor 6 hours in a 5 mg./ liter

solution of indolebutyric acid rooted 30% in sand after 68

.‘

days. Ten percent of the cuntr in this experiment rooted.

\

‘l [0

Greenwool cuttings of . uao'rifera treated by Afan-_4I_ II

U!

I

siev for 24 hours in a 20 mg./ liter solution of indole—

butyric acid rooted 50% in sand after 65 days. In this case

none of the controls rooted.

The results obtained in these experiments are indeed

promising but further work is needed to test possibilities_}

of improving these comdaratively low rooting gercentages.

Studies of tree ages from which cuttings are made, as well

as the rooting response of various parts of the tree, as

pointed out by Thimann & Delisle (1939) should prove to be

promising lines of investigation.

An interesting and unusual characteristic of birch

seeds has been described by Crocher (1930). The seeds,

(I)

when first collected, requir high temperatures (90 de-

grees F.) for germination. If, however, the seed is strat-

ified at 32 - 40 degrees F., the germinating temperature

gradually drops until at the end if several months the

seeds begin to germinate even at these low temperatures.

JYTeLCG

Chromosome number

(-+~

Li (D

s oi chromosome numbers and other reports by var-

go

Jorrensen, 1925; uoodworth, 1929, 1930,3

h

ious autlors (Helms

1931; detzel, 1929; Jaretzky, 1930; Gaiser, 1930-a, l930-b,

193 Stebtins, 1938; nettstein, 1939, adj bleier, 1954)

in

3;

ndicate that the basic genom in the genus Betula = 14.

C)

w c—f.

;)

(unoodmorth (1931) in View pclyploid series in

$13

,_J

this genus suggested an orifiin basic number of 7, altho

no species uitn this haploid number is believed to be extant.

Hanscher (1934) on the basis if interpretation of secondary

association at metaphase I in this and otder genera of the

Betulaceae also sue ested this ultimate basic number.v

Heteroploidy

The comprehensive researches of Joodworth as cited

above indicated an interesting auto- and allopolyploid

3.);

escribed species hybrids and varieties inseries of the

the genus. host cf the menbers are normal diploids (2n = 28,

as g. niera L. and g. lenta L.); two are triploidl (3n = 42,

as p. Jackii Schneider 2‘s. cunila L. x g. lenta L.); four

are tetraploid (4n = 56, as g. oumila 1.); two are pentaploid

(5n = 70, as g. papyrifera Larshall); and three are hexaploid

(6n = 84, as g. lutea hichaux).

0 H)

siecial interest to the tree breeder in this connec-

tion is recognition of the fact that the hexaploid, g. luteg

(yellow birch), is commercially the most important of our

Both are natural hybrids described by taxonomists.

_1nv-

native Species. The production of other polyploids by arti-

ficial meals Lithin t is genus as well as in other eneraf',

..lp—

\—

may thus lead dirictly to superior types for forestry pur-

A study of stomata frequency on leaves of the diploid

niera, pentaploid B. 03yrifera, and hexaoplo id B. lutea

ItD

(as well as Lolvploids in ozher' :enera) to ascertain the

fl:

reliability of such ata to be used as an index of chromo-

some number his reported bv bax, 1938. g. niera (2n) showed

an average of 82 stomata per square millimeter, é. pa;yrifera

h discrep-0(5n) snowed 37, and g. lutea (6n) showed 43. Su

ra and g. lutea(I)

ancies as here aspe r betneen B. oacyrif

were also observed in ether genera. It was concluded by the

author, however, that although such counts cannot be used as

an ab solute index of polyploidy, they may be useful in pre-

liminery srveys, especiNi V of herbarium material.}

Cytological studies of species, varieties, and hybrids.

O

The following summation of meiotic behavior as noted in

several of the species, varieties, and hybrids in this genus

is based entirely on the observations1 of .oodzorth (1929, 1930,

1931).

Di olo ids (2n =28)

Woodworth reported meiosis normal in B. lenta L., g.

nicra L., g. coerulea-(raiiiis Blanchard, é. fontinalis var.

1

Material fixed in Carnoy's fluid, imbedded in nitrocellulose

sectioned 10?N thickness, stained with hai1enhains' iron

haematOX"1in.

NCTQ: :etula, Alnus, and Zorvlus mature their pollen in “e

+1

fall; 0s trva, Cstrvosis, and Jaroinus mature pollen just

prior to shedding in the spring (Woodworth, 1931).

b., B. gendula Roth,CI

H- I)

~ g ,1 0 fl 0 r‘c ‘ ”I" '4‘ - r‘ ,‘\ 0 '

Jioeri (crit. ca 5., c. oaconiCi

é. gopclifolia garag., *. maximowicziana Eegel, and g. utilig l

l

var. oratti Eurk. but very abnormal in s. sedmiatii Hegel,

an oriental species. Asynacsis involving a variable number

of chroaosome pairs nés observed and resulted in frequent

polyspory. Normal gairing, however, has frequently observed

in this ssecics aid the aut:or concluded that a large per-

centage of morpholosically good ,ollen resulted. hybrid‘t‘le

. a“ - -- . V ‘N‘-,r\ J— —‘ A ‘ V A: (-1

origin has SUt355b€1 or b. scniii

Triploid hybrids (3n = 42)

The tricloid hybrid KB. jac:ii Schneiier, resulting fro m

fthe natural chSS‘of diploid 3. lenta L. x tetragloid s.

oumila L. revealed, as right be anticipated, extremely abnor-

mal meiosis. Variable numbers of univalents, bivalents, and

trivalents tere reported tr Iooduorth and in some cases there

was orserred no cairins at all in the ;rophase of division I.

n‘ g h 'y- I“ “mm, 3‘" (v Pr- . N '- 1 ,-. ,\ ' .—~ . . _ '3 .~ . ,‘ _ '1 . .. ‘ *1, .. .

uflfumus He lat,in5 on the soiniles ”as observed in bond leis-

’ons. Polysoory wgsobserved frequently r solting in great

variation in oollen size. Rosendahl (1916) reported the

oollen of this hybrid 25; sterile.

In a study of the comoarative anatomy of tne stems of

B. iackii and its rarents, Cousins (1933) reported an inter-

mediate size and tistribution of vessels, width in cells of

11

compound rays, and will characteristics of bith cells in the

4

set to various other characters tie hybrid

resembled either its oollen parent, g. linta L. or the mat-(T

subcoriata

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“I (WC)

N ‘1 : ‘ ~‘. I r“ v -~ 1, r? :5 a F‘ r (v - 7 .- « ,-. v ‘

a:rg., :- olxila Var. i;¢JJU1lftT3 Hegel, ifl; d. glgyrifera

var. ooruifolia (negel) Fernall mere Std ied oy noodwortn.

6 CV

'3. y-In all asses, o‘coeot the latter, meiosis v.2: normal. Co C.)

ionel rsynegsis of a few hooolo;ons CFromosomes at ii k’nesi:CU

l

was observed in tge oornifolia variety. Some p.m.o's. nere

. -..-4. . . ...:

, not occurrei. In

\ l___J

O H.

5.1.1

'0

O H *._l

(D

:5

on the US$18 0; its very apnormel meiosis, Wes oonoluie; by

Loeitorth to he of hybrid oridin. Parentaje, involving the

(D

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d"

F C.

O O (D

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C)

H O

T he only penteoloiis stoiiei :y woodworth has he white

or ,ager Circa (é. oaoyrifera larsball ) an? the variety

kensioa Henry. Leiosis was observed to be normal in the

variety but slig_tly ab.‘q1;fl hi tue s ecies. abnormality

(ing chromosomes in

’

It is of interest to note tbet the true soeoies, g.

anyrifera, an; only one of its varieties (kensioe) are

(oooifientalis is nexaoloid. in the tasis of such cytolog-

ical evifienoe jg woulo see: that some of the varieties are

fieserving of sjeoi:"

hexagloifi s_ecies (5n = 84)

btuiies o? meiosis in g: lutea Liiiaux revealei soae

abnormality. Ccaaslional univalents a;i trivale ts were

seen at diatinesis accompanied by latying of univalents on

the soinile. Some :efeneratei micricytee .ere poserve’ in

the gollen :roducei.

leiosis in th hexagloids ;. rrossa Sieb. a i Lucc.

anj g. uaovrifera var. iéciientglls (Hook.) barg. W08

reported as norxal.

The genus Petulo trsm tie stanflooint of the cytological

eviience so far accumulatei, aooears to he a group with con-

siierable :rooise of Vieldinp furtgcr valueble evidence as

a contribution to a solution of the questi n of species form-

ation. Prov the evijenze so far available, polyploifly,

especially altopo‘vp omiy seews to have be:n an ifiyortan

vehicle of species for ati‘n is tnis enus.

Jonsiiering fl; tealtn ‘f Iild waterial mnicn is proo-

ably in existence as first generatioo hybriis, segregates of

F2 an? later renerations, backcrosses, etc. it is sursrising

that little or no atte pt las be o TS“€ to select from CH3-

~111—

I-:71 J o

(petula)

Atfl:e*~s, 1?. E). (lfUia) i”rer:ticxi1 TTJthisli Etirefgtrgfi, XAfIIUI+

C 7 pp., 14 plates. b.1016 University Press, London:

Iu.plroj lilford.

Afansiev, Iicrel (1939) Effect of irooln»n+wman acid on

rooting of greonwood cuttings of some eecicwnw forest

BPOOS. JOUP. VUTustry 3 7: SV—Al,

F‘leima', ll. (19%34 Essato‘vnferfinolckyio. ([email protected].)

Tih110“r. Geret. 11: 393-459V

91““ '1, PI. 1’. 3‘ P '-siriri, . :1. (13345?) (39;”U*J101131 'tiifl1e1“s (3f

*le Unitoo FLHLC§. let ed., XXI+~334 oy., 312 tis. to-

3r21~4 ill 'Mxfii CQYQKMIJ, IEM:., Jea'lflvgk 9'”1 01C(“'-

'COusiiis, Cu 3.. (léfifii) EWM; corgwlratiJW3 shrtFW;T of inns stems

of‘ Pedrllez ::7111J“, 11913119 'lerita, 1anri trcz rLlfiritl 3<1t1§£1

1 clii. J. erold Arb. 15: ‘351—55.

Fwocceftql. (1930) Harvesti r , storage, and stratification

of seeds in relation to nursery preoiis,. hlor. Bev. 55;

Ees‘, E. K. (1910) Ibo distribution of self- fiertility in

the flowerin‘ slants. Proc. Amer. Phil. Soc. US: 449-

}aiser, L. 0. (1930-3) 9 lisu of chromosome numbers in

Angiosperms. i’}er...otice 1.9,: 1'11- ? o.O

-------—--—-— (1939-b) A list of chrorosomo numbers in

Angiosperms. Bibliou.. Genet. s: 171-466.

1933) A list of clomosome ru611rs in

nblmo. Bib iorr . reiet. 10: 1 ”-

Barlow, V. L. Fairer E. S. (1941) Textbook of oerdrol-

ogy. 2nd Es. XV-+F4r} no., £94 118. LCGrsw-Hill B 0k

Co., Inc. NeM' York ....

lkeiberxj, H. if. 11. (13;.37—:31) (ii slu‘e“ CM“ the Iiroverun1ce

problem of our 151.031; 11'117,.orte.n.1‘t‘ores t trees, pine, spruce

and birch.) Lcdd. florske Sho'forsoksvesen 6: Befte

20-2.7: 51"101: o

Heimbuwger, C. (1949) Report on Poyler Myoridisstion II.

Forestry Chronicle lo: 14?

1181918 4.1111. n

Dansk.

Jablokov,

10 V31”);

\

—-----—-‘----

nr O

L)

L.Jo

_O_

D-

it”

'71RIJLJ .

Wkere

Birkene pas iedlemose.

I

in the F Ginter-pocif-

* n

1‘r3

C.)

I

Q ‘0

(1

Bot 10: l“9-l 3

(DO)-

M m 1.1 J D

C+

NJ

1'!

A a

b

qr»es

eh

TU

(n

uiJn:

'J

3‘s

{,3

v

1

C‘-

JO

JP

10 o O; D 3

w

3 j

1...!

l)

"U

H u b (’1'

0

“is . s. . .

Jentys-Sze r>ws, J. (193U) (Biometric91 studies on the

col ect ve sgecies E. aloe. L. 11. The possibility of

hybrioi "tion between specie B. Verrucooe Ehrh. and R.

nuoesceos Lhrb.) Rosprssy i sgvsuosdanfa, Inst. Duos?—

czy as67 fans to'Tcn, V»rszswa No. 4 : Sor. A.: pp. 84.

(Cited from Plant Breed. 701. IX, 19.39).

Johnson, L. P. V. 193$) A descriptive list of nntursl sad

srtificial interspocif1c yorios in North American forest-

t~90 genera. 99996. JOUP. Res. 17: 411-444.

-ll2-

Larsen, C. S.. J ‘ 3

indijvioiulls -11 i()res1dT~

I113 Of snecjes firings; our”:

8

L a Q ,J

U O V .1 » 1‘ If. . ska [VSL'(‘L‘~I' ; E)g- L40:32.

‘ T ”'V‘ 1" '~ ‘ 1 - ‘o P‘ ‘H “VA." "3‘" ' ‘f ‘\

See also: A 5. vet. 1L-. Joil. 1e11o00L, CLtiliven,

1 ° _ ... fivr '- i . 1-. 1

Peprint yy. ., 13:7.) (Cited irom Plrnt Preev.

{11111 S t T! . 11701 17

.qn_oe, P. F. 19) The Kerton cetqlo118. A list of the

c romosoue n lerrls of species of British FloweringBantJ.

:1: 011v PilbrtOl . ()1sz 1-51 .

LorQen+PQlep, H' (1915) WeitrHQe ZN? Kenntnis des Fermen-

1iPej-Ses €99 SNTWCl9Pt ?Cffllfi 91h? L. ’qvicir], 12115. (Cf.

Lens: Skovf, Tidsgkp.‘fif“ggg, 19 7.)

Pearson, 3' 8° 9 3:0W3, H. P. (193?) Commercial timbers of

India. tsleutt Government of India Central Publication

Branch.

Pendel, A. (1940) kennel of cultivatei trees and shrubs.

2nd Ed., XXi-9 5 pp. LACvillon Co., Len York.

’osendah1,C. O. (191U) Mervtions on Petals in Kinneso-

ts nith soeciel reference to some nature uyorids. Linn.

Bot. Studies 4 (4): 443-459.

Sex, H. J. .(1938) ”Be relstion between stomate counts and

chromosome number. Jour. Arn. Art. 19: 437-441.

Schreiner, E. J. (1937) ImprOchent of forest trees. .8.

D.fi . Veerbook,193 : 241-1279.

Segreinej, E. J, (19.38) Lubit of forest trees.

Ciron. cotanicq 4: 495-494.

Smith, E. C. & Lie‘ools, 3. (1941

trees. JOUP . Alfl. Pad}. 32: :

SprenQe:s, (1937) Sen en ands

(Iotes on the cultivation of b

Heidemsat son. 49: 191—19U,

Stebbins, G. L. Jr. (19"U) CytOIO“ical crwarecteristics es-

socisted with the diffelent “rowtt stits in the dicoty-

ledons. Amer. Jour. Bot. 25 (3): 189-198.

Thirnsnn, K. V. & Delisle, A. L. (1939) The veQetAtive prep-

egetion of difflicult r1Isnts. UOHP. Arn. Arb. 20: 116-

106.

.snscer J. H. (1934) Pssic01110mosone number of the higher

plants. I.Iew Phytologgist 53: 101 f.

Wettstein, 9. Von (1929) Zur Technik der kunstlichen Kreu—

sung bei Veiden (Solis). Lfichter 1: 125~ BU.

----------------- (1930) Lie Lrenzu1Qsmetnode und die Pg-

scbreibunQ von Bl-festorden oei POpulus. Pflsnrenzucht-

EilOijllg

E>eeies hybrids in forest

A

over de teelt ven berken.

_rch trees.) Tijdschr. Leo.DJ

IUIS (I?eilu3 A) 1U: 5E”7— 6‘?, .

---------- ------- & PPOpacn, H. (193”) Sichtuni:sgrljeit Qur

BizY1nzicbtu1Q (Peconnaissance work in birch brreedinQ.)

Zflchter 11: 379-UO. (Cited from Ilsnt Breed. EAstr. ”01.

x, 1940.)

Wetzel, G. (1929) Chromosomenstudien bei den rstles. Bot.

Arch. 25: 25 ~2U3.

IPaiert (1904) Betulscese. Des Pflsnzenreich IV.

61: 94.

, H. end Anton, L. (1933)Fitudien13oer Petule alba L.

1m Ansc‘rvluss an Ia-‘lorQent‘rieler 11nd morqrsson. (studies on

B. aloe L. with reference to the torL of Iowzentnole“ and

iunnnrsson.) Beitr. ‘ 101. Pfl. 21: 255- 99.

-1;13_

floodwopth R. H. (1929) ytoloical studies in the Petu-

la eee I. Betuls. Wot. G-Z b7: 331-363.

------------ ~—- (1930) T1010ioa'il studies on the Betn—

lecgee IV. Retule, ngSinus, Metrva, OStPJOpSlS. _Eot.

G53. 90: lOb-IIS.

--------------- (1931) Polyploidy in the Retulscese. Jour.

Arn. Arb. 2: 205-217.

Yanchevsky, K. (1904) On the peculiazities of the germin-

ation of the seeds of Asoen and oi some species of YiIIOW.

(In Russian.) Proc. I“*e*1il r0orest. Inst. ll: 2”9-74.

LiszsirUas mos CITED

(Petula)

bianca, S. (1933) New principles in he collection and

4

.1.

b

preservation of forest seeds. Int. Rev. Agric. Ftome

508-17

Dena, S. T. (1909) Paper birch in the northeast. U.S. For.

‘erv.CiIc.163.

Fernald, I L. (1902) Relationships of some American and.

old morld birches. Amer. Jour. Sci. 14: 167-194

Ishikswa, m. ( €15) A list of the number of chromosomes.

Hot. Rag. Tokyo 30, No. Boo: 404-440.

Jack, F. G. (1895) Hybrid bircres. Tardb‘n & Forest 8:

243- 244.

’L tZ7el, G. (1927) ChW0uosoncn Lnlen bei den Ferles.

(VorlHufie Litteeilung.) Ber. Dent. Bot Gesell. 45: 251-

50 o

-—----—-- (.1./(3

(VOll'lgdfive 1::

l4.

‘Ioodworth, R. H. (I929—a) Eo‘to99039nesis snc nolyembrvony

”Inomosoverzablen bei den Fegeles.o) e

itteilnng.) Ber. Lent. Pot. Gesell. 46: 212—

\(t

LuRoi) Sureng. Science (n. s. ) 70:19p-in AJJNIS 1921939

193.

----_-----_-—-—- (1929-b0 Cytolo*ical stud-

wilece9£>. II. ICOTQThls91ml Alnusq 130t. tc.l. 85; (JbO-

39(5) .

--------------- - (1950) CytOlOSic9l studies in the Petnlac-

eee. III. Parthenogenesis and nolvemorvonr 1n Finns

rm )se. Bot. Gaz. 89: 402-409....—ALE;—

‘ I ,‘\ ' V'Y-I ‘.

I‘A.'_:.'."L!3lbh_4 A. PI‘QUH

Beech Family

‘he beech family includes over 600 species in 6 fenera:(

l

\—J

Quercus L. (oak), Facus L. (beech), Castanea giller (chestnut),

.3

CastanOpsis Spach (ctinquapin), Lithocarvus Blume (tantark),l

and NOthOfQLUS Blume (antarctic beech). The family is found

thruout the temperate and subtrOpical regions of both hemi-

spheres but is most characteristic of the forests in the north

temperate zone.

With the exception of Nothofagus, whose 17 species are

restricted to the southern hemisphere, all genera are repre-

sented in fiorth America.

The economic significance of the Fawaceae rests mainly

upon the production of valuable timbers, especially oak, which

enjoys a world-wide reputation for strength, durability, and

all around usefulness. The family in addition produces other

products of great importance to man including cork, tanin,

and nutritious nuts. It has been claimed that no other family

embracing dicotyledonous trees in the north temperate zone

I

p

rivals the Panaceee in their service to man. (Pearson 6 Brown,

1932; Rehder, 1940; Barlow & harrar, 1941).

Que rcu s 1;. (oak)

The oak genus includes about 200 arborescent, rarely shrub-

by, deciduous or evergreen species in the temperate regions of

the northern hemiSphere and in the trOpics at high altitudes

south to Colombia in America and to the Relay penninsula in

-115-

Asia.

_uercus is divided into three suogenera: (l) gyclobalan-

oasis Prantl; (2) Erythrobalanus Space (red or black aka);

asi (3) Lepiiobalanus Enilicher (White oaks). The latter is

divided into 6 sections: Zerris Loudo: éEEEE Eeichb., lléfi

Loud., Gallifera Spach, Pobur Feichenb., and Erinus Loud.

Cyclobalanopsis is not represented in the flora of the United

U)

tates but the red ani unite oaks are represented by 75 to 80

'ecies, about 60 of which reach tree size.(n

'O

The norli-siie reputation of oak timber has been previous-

ly noted; but it is also a matter of significance that the

genus is gregarious ii the north temperate zone, forming ex-

tensive forests in regions there civilization attains to a

(D

U)

C D (D

O {‘3

...H;

r‘

‘e of specialization. In the United Stat

furnish more native timber annually than any other related

group of troaileaved trees, being surpassed only by the coni-

fers, which occupy first place in this respect. The principle

native timber producers of the red oak group include: '3.

bor alis Lichaux (red oak) ani its variety 3. b. maxima (Kar-(D

shall) Ashe; g. velutina Lamarck (black oak); i. coccinea

iarshall (scarlet oak), ani several others. In the white

oak group, 3. alga L. ani i“s varieties are said to furnish

nearly three fourths of the timber narseted as "white oak".

Other important producers of this group include: ‘g. macrocarpa

Kichaux (bur oak) and 44 grlpug L. ( swamp‘chestnut oak).

The two white oaks 3. robur L. (English oak) anl J.

etraea (hatuschka) Lieblein (durmast oak) and their numerous

~116-

varieties are the important oak timber producers of ;Le sritish

Isles and southern Europe.

The cork industry, centerei in Sgain, southern France

and Algiers, is dependent upon its source of raw material

from the outer bark of 3. subs; and the variety 3. 8.

oc iientalis (Gay) Arcangeli.

Tannin, extracted from the bark and leaf galls Oi many

L):

native an. exotic oaks is used for tanning and medicinal

purposes. The acorns of certain species1 are used to a

certain extent as human food but more extensively as mast,

i.e., hOg food.

The genus is tf little importance in the orient although

small quantities of Jajinese oak have come into the world

market from time to time. Thirty-seven species of guercus

attain tree size in british India. Cf these only 6 are

classed as low-ranking commercial timber trees. (rearson &

Brown, 1932).

Because of their sturdiness, longevity, and the brilliant

autumn coloring of their leaves numerous species are widely

used as ornamentals.

The extreme variability exhibited by many species within

the cenus furnishes as yet unsolved taxonomic difficulties.

The description of many known or susgected natural hybrids as

well as numerous varieties and forms of doubtful rank has

abundantly contributed to the confusion.

l

Quercus cornea ? of China is reputedly an excellent quality

nut for human consumption, Fairchild (l959).

-117—

ARTIFIJIAL adiajTlOS F?;l ”l_D Sftjh

Natural intersgecific hybridization in us rcus, based

largely upon taxonomic evidenCe,.is aggarently qaite common

and explains in oart the characteristic variaoilitr and in u:-

gradat ion of various characters between species which has made

this genus the most complex of taxonomic puzzles. Natural

hybridization aogears to be limited to members of the same

subgenus. In the case of the white oak subgenus (Legidobal-

anus) intersectzcna1 crossing has been noted but for the

most part is limited to species of the section frinus (Tre-

lease, 1924; Johnson, 1939). Artificial inter sub-3enus

O

crosses (Hrytirocalanus x Leo iddobalanus) have, honever, been

reported by Pjatnitskii (1939). Johnson (1939) listed about

75 natural hybrids in the genus.

J

It is beyond the scope of this paper to review in detail

the abundant taxonomic literature relating to natural hybrid—

e of the sub-(D

ization in this genus. Eeferences to this pha

ed in the "literature not cited" list, p.|27;L.‘J

ject are inclu

As pointed out by Johnson (1939), it should be borne in mind

that C+

he majoritv of natural hybrids in the genus have been

presumed by taxonomists on a purely morphological basis with-

out experimental evidence as to parentage. Although such

hybrids must be treated tith some caution, the considerable

degree of intergradation between sgecies would favor the

supposition that hybridization does occur.

Selection of superior forms from wild stocks in the

genus has not, aJJarently, received attention in the UnitedL

States but considerable interest in race identification and

~118-

pro eny testing LES beei tarrie d on by zuropean workers.

poerman (1932) reported the testing of 90 samples of acorns

collected from various classes of mother trees in Denmark.

The material was collected and somn in 1911 and observations

were made periodically until 1929. Cf special interest was

his emchasis of the jenetic axiom expressed in his weneral

conclusion: "... bad formation in a mother tree does not

necessarily lead to inferior :roeny

oirner (1933) in Germany stu iiei inheritance of e;icormic

branch (tranches arising from dormant buds) formation in the

oaks. From data collected on the origin of oak seed sown

over a period of 50 years in various for=st plantations, he

concluded that epicormic branch for nati‘:n has a character

thich varied according to racial types or varieties of the

several oak s;ecies investigated. The author recommended

that future planting material for use in Germany saould be

obtained from the desiratle races and varieties he had

selected.

According to Staf and Teerink (1936) 3. robur has been

‘ation Wltfl regard to race iso-he subject of long inves

ins of this speciesmlation in murope and several distinct str

are aid to have been identif‘ ied. Sipkes (1938) in Holland

and various other juropean and British workers (hatthai, 1922;

Hauch, 1909; and Day, 1934) have contributed to the problem

of seed provenance and race isolation in the genus.

-3C§TFCLhD :E‘DI- %;J ELLEJTICN

Artificial hybridization studies in .uercus have been

-119-

.1

reported by a numzer of workers. Cf special interest to the

l)

forest a' neticist is the fact that hlotzsch (1854), first of

the tree h bridizers, success ully hybridized i° robur L. x

0

age ‘.“VI—J (

D

I.J

}._)

iJ- ‘3

[..J

O—

o:a Salish. and other species in the genus

'52; e 1 TL 0

In 1909, base (1918-3, 1918-b, 191;, 1;27) in the

States, success;ully crossed

J..1. _ \ A1 _ _ . _' ‘3' 7‘ 1 3‘ O f! I “ _ o ‘_‘ " .. _ I“. _ .

“alter and noted udrkbd hybrid Visor in the pr03eny.

( _

W

,_J

\0

1 rate

Yarnell

'33) measured diameter at breast height (4.5 ft.) of two

of the hybrids 24 years later and found them to be 13.4 and

S14.8 inches respectively. Eleven 3. virfiniala parentr

the hybrids thich tere 21 years old averaged only 5.8 inches

in dia:eter. Twenty 4. lvrata parents, also 21 years

averaged 5.97 inches. The author concluded that variations

in site or "he 3 year start of the hybrids could not account

for this marked differince in diameter and that it might

logically be attributed to heterosis in the hybrids.

Yaroell also studies seere ation of leaf edaracters as

to size, shape, nature of epidermis and several fruit char-

acters as expressed in the F generation of the above cross

2

and concluded that in most cases only 1 or 2 factors were

involved in the determination of each of the characters. he

pointed out that such a condition increased the chances

securing desirable combinations of characters in second

generation proEeny.

With reference to the s

Schmidt (1930) reported that the type of axis, certain

f...)

An intersectional cross (Erinus x 11ex)in the white

subgenus (Lepidobalanus).

~120—

tudy of heritable characters,

'I 7

0 a5;

1

crown ani 1e;f trees, and grocaely percentage of summer wood,

resista.ce to nrought, frost, disease, anl other characters

have tegn demonstrated as heritable characters among members

of the oak genus studied in Germany.

Following the controlled hybridization tor; of hlotzsch

ani Ness there has, a);arently, been no reported effort of

attempts to conduct furtfier hybridization nor: in the genus

' I'\ "\ l . 1 A, J.

until 1539. In that year Sonner (1939) reported successful

duplication of Ness' A. ”irniniana X lvrata cross and a suc-

cessful 3. virviniana x‘g. serrata (?) cross. rjatnitskii

(193)) in Russia successfuLly crossed 3° robur with 4-

. y. - - 2 o 1- ‘l

borealis Waxima (aarsn.) Ashe , 3. porealis :icnaux , J-

macranthera Fisch. and Jay. (and also the reciprocal), and

'I z r . .~ - . ' 5.1 ‘. [-1 ""1 .- . 4“<f‘ -:‘ ‘1‘ \ —‘l- o 2 *

macrocarya Michaux. lne cross macran.hera x corealis and

1

. 2 a

e lis raXima mere also successful.C)

Hmacrantuera x b' '\

The four hybrid or ogerles resulting; "rozn the inter sub-

genus crosses showed the most marked heterosis and of the

three, the most promising has sail to be the 3. mac ant era x

borea_is noxima eroup. The same worker (1934), as a result

of self-fertility studies in the oaks, concluded that self-

nollination studies gave great oronise of yielding new and

valuable forms from the apcarently heterozygous oak seecies.

1 , . ., ,Johnson (1939) listed only the artificial hybrids of hlotzsch

an_ Tess but there are doubtless numerous unreported cases ofI

w

artificial hybridization in the genus.

lntersuh-genus crosses (Lepidobalanus x zLJLsrcealanug, Hue

first reported in the genus.

-121-

. . 11 T , 1-. . . " , ,. , . 4.“: n r , f‘ . .« :. .

8105:.) {.0 31-13 .-. X g. 1"»)...‘tar L. Cell”; must; .1 J; C5188 OI LIthXCNlfi.

:1 3, r? ‘. r I‘ - \ -: ‘-_ g . I \ :: v ,I‘ r ,. ." .. , .C‘ . .- n, ;. - p;

as 134.catei t; the celayei 2atu1t, oi accrns on the iemale

, ..-‘.‘ y. _. , “ , - _ -4,” 4- ,. 4 ...A , - —. ‘ ,. _, ‘ fl . - - -. i 1

parent. 1t VHHJ.JLS note 1 unit the hyorio.2mnnn1s apg-arecz

‘ L "' r A. _ — 4- J ' ‘ ..

t0 e8 somehzlat .L'il Cf snail 211 St, -ka/(TCJ cu "L'ltl'OL-€u 11’]-

s ‘ u. ' ‘ -— \ \

trasr“01fic Jollcnat'-n on t_e feriLe JlEHt.

The benus is monoecious, te flower dichoiamous and

usuctly taruec ey such extreme grotaniry that isolat trees

.2 r" " L ‘ '. ‘ u ,. -‘ ~11 ' V V ’3 f‘ 8' .0 . _‘_ J_ :1 _ ' J— 1hr I ';-7

uU not eioiuce iruit. lnvescl;atiens by: jatni ts8.11 (1534),

nettstein (1935), an: east (1940) have estaolished self-

fertilitj for the genus as iar as known. ochreiner (1938),

A ‘I' '\ ’ 1‘ '! . _" "‘ 3 v '. “‘0' ‘ . “ ' ’ ; +.’ ' + ‘ " ':‘

Uh LaL/L’: b.1b18 0.1 \Jl CllLAlerff St«Ll-.l:S, TEQOI en Luau bk).-.e

Renters o: the genus ruercus have seen include: in the

”difficult group” of slants to root from stea cuttings. (The

recent usrk of Thimann and Delisle (1939), however, aspear

to contrioute to its grompt removal from that category. These

workers mere successful in obtaining 82% roo ing of auxin-

treated cuttings of tuercus hor alis and 22% rooting in un-\_.

treated controls. Cuttings were made by tLese horaers in

geLruary from 2-3 year old tooi from t-e -ssal parts of 4

year 011 trees. The Cases of phe cuttings tere immersed l-2

(\ A“

cm. does in a 400 mg./liter concentration or injOle-3-acetic

acid ftr 24 hours. Jontrols mere oLaced in tater tor a like

placed in ,rooagatin: ooxes fitted Lita a sic e

closed. The soL F (J

'-

¥1

(

3

(T)

(C

"1

H '_.l

‘<‘.

r It

(D

U (7*

pd

(h

S. *‘i

‘__J

A ‘0

These torlers e Dhasizei thai.t&n3:2ost imeortant single

-1

factor involved in rooting ,uercts an; other ciff

cuttinrs is are of the tree from hhléfl cuttines

T .‘ ‘ 'Z A 7‘31. *0 f q-v 1‘ 3 w-~( C1" 3;)C'yr,j-\_" Qfiwn

rEEb /- . ELQIL‘J (J Q,-"le CirL .I.\.-'UU ._l.vl.JlJ. LA-LLe U._.A‘\Je

cuttings are coscioie without ca ,letely aestro

re 313.518 0ST),A J- n .-2 . c, ', a; e -, 4. .O . -‘

slants lr‘Oul miijz.‘ L..k.' C‘tlb+ll’l~.b

\ r. x" -‘ ' a“\ I | ' ,

-.J .1.]. v1.1an

Chro; some numser

'\ 4 ~ ‘1‘, " ,n ‘ r ' ,' r' ‘~ V T v r" A

:esec on the chroiosome counts of joey, l9

‘-‘ _ ' P _ ,- r—V‘r ‘vafi --.!“ .

1929, 1930; sex, 1;3e; caietzny, 1:3»; flgflOll,

1940 the haoloii CSIOZOSOMB number in the genus

'orted in('2

Various other counts have teen re.

(2n = 8, Cosens, 1912; n = 11, netzel, 1920; 2n

1930; 2n = 12, Aufderheije, 1931) out sharp (19

Ho

finII

‘29;

34)

8; ti;

"1“,. a

bull

(33"!-

LA“;

mpu

Friesner,

Luf-

field (1940) concluded that the {enus was homooloii With a

r,

pI

C—f—

H.

< H.

L/.

t,

somatic number of 24.

F).

a basic number of 6 for the genus. he tel

number of 12 chromosomes osservei in to: variou

studied could be civideo into two groups of 6 oase’

L Eee also li t of chromosome numbers by Caiser,s s

and taude, 1939.

-123-

C:ie (1937), however, suggested

eved the reduced

3

morohtloer. Iie also resort e; seconiary association of mem-

bers of the the groups at metaphase I and 11.

K)CytolOnical studies of s ecies anl hybriis

The majority of cytological work conducted thus far in\

the genus has been executed primarily for c roxosome number

'7 ‘L "’ ° '3 v" P“ 1 :3 7' ~ ' '1 '1‘; "1 r~ '3 ‘r‘ I" v~'~ r1 '5} ,- fl : ~ ,-' in. 3 r') ' r‘

38 Ltrmlne~t .LLJD, Ll-1JCL’1 C‘l "LJ._L'J&1 L-C;..|:I LCC.1 ‘-_L'~_‘Ile if] SOLILCZt-LD tlSDueO

The majority of the workers, however, have noted that size,

shape anl number of chromosomes is remarxatly cons ant from-..-

seecies to soecies. Chromosome studies based on counts at

$0

meiosis save ls o revealed exceptional regularity in the

meiotic divisions.

Sax (1930) estimated pollen grain sterility in 25 species

and hybrids of the genus ani found it to vary from 3 - 10%,

except in one excegtional specimen of 3. dentaia Thunb. which

snowed 80w sterility. Another individual of the same sgecies

showerl' only Sfl. The five hybrids included in the study nowed

no significant difference in sterility from that found in the

e pollen ,rain size has also computed for

eacn soeCies, and this Character also showed remarkable uni-

}—+J

formity (except in the case the exceptional 3. dentata

mentioned above) Wlich had notably larger pollen grains.

The genus guercus, in view of the dearth of experimental

q

genetic and cytological data so far collected in it, appears

to be a fertile field for basic cytogenetic research. fiith an

increasing perfection of vegetative erooag atitn, hybridization,

and chromosome doubling technLuse this already too-ranliing

forest tree ::nus may make even more desirable contributionsk.

to the constantly increasing debands of modern culture.

~124-

ll‘"?xfb"

siev/31.13)(

m7""

(JI .LJ-‘A

llifderreide, .1. (1931) CE-nowro OzW‘YUULDGPS in.PVfijxs gn~nl-

ifolia and ,rsrcns Virginianc. Antler Univ Lot. Stud.

2: Eager LO. L‘: 45-52.

9 rner, 9. (1953) erer lesser‘eiserhildung, his} ;wnassen

11ml E1036}nx/Hlfnolz zucita (Q3 tlcthnFMJtiOn 0i e;pjicormi

*ranches, Oil laces and the production of large oak tim-

ber.) Ltscb. Forstw. 5: 541—544.

Conrer, A. B (1939) 52nd Annual repart of tie Boxes Agric

Station.D‘p-

304

C

Cosens, A. (1913) A COHtPifigtion to the wohyVolOJy and bi-

OlOJY 0f ins Got 63118- PPans. 31nd. Inst. 9: 297-5sl.

Day, fi’ 9. (193') T99 Pel'LiUB 1Det‘rfeen disease and the con-

Stit”tion and “““ironmeHt 09 t'6 tree. J. a. i;ric. Soc.

95: 54-72.

Ullffield, Jo .1. (ll—440) C:'I'O:::OSOT:’.O 001131133 111 ”9190118. 53.11191“

'3 301:. 2'7 (s): 7:27-53.-—

East, E- 5- (1990) The distributior of self—st rilitj in

the flOWBPin; plant”. Proc. finer. Phil. Soc. 62: 449-

51s.

st_rchild, David (1539 The world was my SGPdGn. XIVfi-

494 IM?- 33953. Sorfisni r's. lien Ytnfli.

Fouarre, J. (1939) Note su_r 1a cntyocinESe crez 1ees cliones

pedoncule et TOJVPGo (3 note on keryokine-is in pedun-

culate oak and sessile 0k.) Hull. Inst. awror. sembloux

8: 111-113.

Friesner, R. C. (1930) Chromosome numbers in ten species of

"iuercus, wit- some rem r1:s on tie contrilmtions of cytol-

05y to ta monogy. Lutler Univ. Bot. Studies 1 (Papers 6-7):

77—105.

Gaiser, L. 0. (l950-a) A list of chromosome numbers in

‘ioene““s. 11. Si liegr. Genet. o: 171-4oo.

-----------—- (l93x-b) A list Of chromosome numbers in

Angiosperms. III. ienetica 12:101-200.

------------- (l953) A list of crnomo"ome numbers in Angio-

sperms. IV. Biblicar. venet. 10: 105-236.

Ghimpu, V. (1529) Eur les Chrome some 3 de ouelque ctenes.

Pev. de Bot. Ancl. et d'A3ric. Colon. pp. 176-179.

---------- ( 950) Vecherches CthlOgicues sur leg genres;

HOPGGQEL_A?H019, ice'ce o, Vitls 6t 3ncrcus, APCP. Angt,

Ticros. 20: 9135—2 30

Farlow, 3. N. “ Harrar, OE. b. (1941-1)lextaook of dendrol-

2y. 2n} hd., XV1'542 pp., 254 Ill Icfiraw-hill Rook

00., Inc. 13w Yor & London.

Hauch, L. A. (l999) Evblichkeit bei buche und eicLe. Cont-

bl. fiesam. lorstw. 35: 355-343,

H063” 15. (1929) (fillflellerlTV‘TWWNDTnCllenlfjnnwvis robnp I”

o; J. sessiliflora iartyn. “op. Tide. 9038H~1;6n 40:

[11:409’

Jeret"'“' Tl. (1930) Jar mytolo;ie der Fagales. Plants.

Arch. Kiss. Bot. 10; 120-157.

-125-

181, 0

1.01141“

17:

WMBIM

Eelwi

’i Y“

nscni, A. V.

srtifici:

1h

1]](1.L .

J.

C U

.,—o ‘

.15;

sPGC

a

I). J-FCQ.

Che HuinfunQ de

es. iSthLCL. Q

Vein tr. 12:

Lerton ostsloQue.

species of fiIitis

d 01. 3s; 1—51._Q

(1257) I: LIQ“~)e

hioridos do Qencr0

5ndh;L

I?“:ukflf‘0638 (.19. 1).]

Ilow i

Forstwi

A '\

I 9

A23

UL

rv ,-' r".‘ ‘_ )o I]

K. I - I 1 '9; .

M

f1

Po

\-

,

qM

5““;

p

S 1

s.)

ervieos

(1,16-9

Jon_n

"C)

{—9 V) '

_Uerou

C108 {3

Oak.

1" ...

rJ OLLI‘ 0

ed“or

hybrid

f “stl01 .3 __

x {Inf-1 dfs‘rl

,

mflmiir.-.-q

r

;\J

O

J

fi J

.__1

f.

..-)

E

P."U

0

Htlmh

U)

(T)

1L

f‘netursl

411—444

Q do?

Preuss

O

148. o

' \1

I u

I23.)

list 0? tbs

onerir=I...)

ices INEl-

American

P frin-

{43 1'1 (3.11 391‘ OV-

5—419,

f‘

(CytoloQ-

tbu(JV!

(J¢‘ 11.8

r. a 0 I‘m 1‘"; 1

7A

en 1 mes

. kOMfliSSi

P:ports wi

p.

e1 timbers

1 Putlioet

Lesnoe

Plant

11.

-.

1):).

A..~

“rsnoh.

Pjs.nitkkii, b. 9. (1934) 'E‘Yeerinents in Sglf-pollinst

of'iJ pit, xuzor srni‘QICrous. :kcta Irmfis. Rot. fxyui. go

1I.S.u4??. 4—'_~§97'— 510.

--------~---—-—--- (1939) (oviniczs.ion in oaks.)

1107 aistvo (borestrV) No. 7: 36-45. (Cited from

greed. Abstr. J01. X, 1940.)

Behror, A. (1949) Assisi of cultivstsfi trees and shrubs.

Bud Ed,, .735 so. T*(fiiillsn Co., INII'YOIK.

Sex, 9. J. (193C) CTICIO omo nurters in Qn_ercns. Jour.

Arn. Arb. 11: 29’—295.

Schmidt, V. (1939) Forstlioto Ffils nzenzfichturg (B_oedi

forest plants.) be? ”ontel 2: 189-199. '

Qarreirer, E. J. (1939) Besesrch in forest 1venetin it

oetresstorn borest Exp. its1ion. Ber. Proc. 29th.m1n

th. horthern not 9r. Ass., Isss.: 12-14 Septowher;

59-58.

-~--—---------- & Lquisli, J. a. (1942) Metsxenia in an

osk species cross. Jon» Eerefi. 33 3): 97-98.

Sherp, L. H. (19"') Introduction to oytolon 3rd Fd.,

XIV 567 pp. 233 ii 8. ECErww-Ulll WOOL 30., Ito. N

‘for” E: ififlldffl1.

Siqflces, 1?. (l?"b‘) 'Ri‘ien. (”WA s.) TiQyflsol “. lkyi. Ihxi’

E 9307?. FT): JJJL—l.4.

~126—

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TP- — 7‘ r- (I. \r‘.

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T“ " r70*

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all. ;0~'o; 'Ot. Cllw 9: 19-15.7.“ f h '1 ' :‘Nf‘\ r- ‘ _ 1 W '

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f“ 1 "7 —,‘ - (7- "7”"

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'JT‘C‘ E OI'CL)11 U: ng-J.

~1nn1nu a ( <n7\ nm-uw wan -m in. a~ - r .V +LL, A. 4‘ u; unh;_suunvlven {per Clo mlvtscrnftllnrn

I‘pf’r‘114‘l Y 1' “I" '5 ‘v' 17- ? J‘ l] , (I. ' 'rw 7 ("- 3 ' ‘,-

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pentul. mosar. f033ua. 4,; (fl7)-149.'V\ O .‘I' '1 \. "'\ 1 1.. 0 1

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333 c. _u* ex 'slanu rgnc, i; g-7§_

wry. 1 ,‘ - - - 4 ‘

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11:53:" I hr; '37. 1, 3'0 ("-g, ‘. ' Q n '1 " A J- 1 —- - 3‘ '1' - A -. --. V.) J J- . (J- (..L J4. l Lv‘ L’CIL 1..ng .' . k)t.; (4;) Ifulz,.;-k1 $1131). ‘\,I,LS

.- q < 0 1m r‘ r;-

32“ ~01. 1’1’00. (lra‘lo) l: on.

_, “I'm-d A ‘~~-‘ -' ,1! ‘ -' "" - -_’ '-———c————~—u—— (L..L" ) :4; ‘ dl- r.:'L(\' C'{‘1‘,/ .71” f‘er,‘\;-’1J_Eji'}: :‘Jvi' {;,\11

I film) 3 I“. 3'1?) ) rr~ (3,3. "U ’

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law» , In"7 (lu.7 L; ? To” 0:. Viv ‘ - 4- ’1 ' ‘1' u '1 .. \ . g 1_ . —‘ .

asuern (iOMvhlu ?1H “P twe VHPIRUlJFt

:1' ~,‘ ,, °*nr >.r.. ..v ___‘1 3 T4, ‘! . J.‘-3..:l(3-_ - 1 I‘d ~11: ‘ .

l

Fixx i, .\, (17177) LI 8 rvlnc \w‘ric4x‘ e ’"hv- _ iEwirifii 0 hurt?-

301 01 ” ‘KHWNIS r0w~*”' (imwc X“K¢i€ii€8 CdTEjflxrids of

f, anw‘.) (jlteJ—THHW Pl€“u ‘rasd. VUl. V, 193%.)

”1n1, ?. {*"“d) ”b;? 616 I=C-Eo:nw“sa?nft plhfi? Weiss-

iru11“czx lifivw} c k ‘1:'101'“ *4\"3A ‘ 13r7_-*rip'iriiJEL'WKIiss) . (Cm: tE1e‘ o L o . ~. T \

progelj of “ gycgl “c u 1.2 vJa .ueu 0'» \_. TOU“”

ql“i-Ii?ifii"evviso) 791*. ‘ot. :\1. 55: x15I-bfi.

ficJIS, 3?. (1513477 ‘Vii+““” "e (“31"fiesv3‘5-v*n C (30Y*' UFJ‘P V3 SL39-

q1¢ifln~¢ Lvlwsbnry “Vnmfica (EFe nr‘vc) c: 4p4-ZZU.

'qu{57~, 1,. .1. (l JKQ i”'e <)"' ill IIWNWH1%1:. “1’1ficy d «193:0

*‘Ochauqel .usrisirl*orln;jii ”erkmwni 1‘74; $33 ”To

TT¢ENL, 1n. (1‘”‘9) Crl *3 e .7nfl'“""v‘ri" “01“18 r ctnxeaxl ”;19""v F

IWfi*¢? 7}. Pr( —. SV“U‘ilijthPfi.Li%Pt:“l. 1C)t. “WK3QGI£:TKZ?S__

217:??7. (Ififis; & hngfisl swmwrry.)

¥ollfck, &. (109%) A recent Cinnm\crg 6? Ed.rid 0¢ks on

ctnten Islfind. “all. infirej wt. Club 1?; 303-399.

---------—- (131$) Tbo (tovy Cf t=0 antrnn 03k. Sci.

; gw‘ 7’0. 4vfi9, f‘“-/33, ‘432.

}”CL, J. C. (195:) ”9rd“ 0? Eylrid 09k extcbdec. Jon”.

forestry 3”: 701.

Kline, .. R. (171*) ‘ rare :‘fixrid 0" *Ylllrn1871VOniq.

FOPCSt Leeves 16: 120—91.

Iachngal, L. 1. (1307) L,.11(129tion of tke oaks. Sci.

«MGP. Q‘p. 03 ( 69L): ZulCE—leed.

-—-—---—--————-— (-997) Tybridififlfion Of the oaks. Sci.

AnsP. ..g;. ;3 (lCBQ); 2LfHTS—Quloo.

.oe‘zy, I. U. (lJWW) chbril.twnL f) the 11?: listory of

:1]_F=r7t S ; t? (3 'Tr21‘1;1“01*1 (Jai‘: i-*' (3C)“‘(;(?t3j.()11 j_a=t;‘1 ' F 3°: q Tri()1w '1rlci

hybriiisn. nerd. Lit. Mei. 31119. Prom. 53° 33-24

Nueller, C. V. (1936) Studies in t}e O“V9 of t*e monztfiirs

o.“ 3:03"+}20°s1:.r‘1m EOJUQO. me .‘1'-.. EXT‘E‘J. 1'7: 100—179.

P9Pk<_, “. Y. (193V) r9'0 nwtwru hymrifis. Indlan ‘Orostcr

CS: fibE-Lfi‘.

PCFGer, A. (1“01) Fo+gs on Pybrids of "~~nts ilic’foli:.

?*Od0ra 5: 137-140.

.cuer, L. T. (1909) fibrous L093“ L L;%_id 0%“ Plant

'0316 l&: 190—20].

Schentz, 0. L. (1335) AH odd yurlfl 09%. RP. Potflv 40:

3—5.

861%“ (liQS) ”raga d1) Lie en, .13 sie farhqen. Ztsvhr.

Focst 1. Jr ow. US. 39 —531.

{Fidvx)”t}1 }. ‘2. (lCVLZ) Ngilrjxi 091(8. “Imyn. Ifi01u33trfiz 27}

d r‘fll»

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6 - 'J.

Treleuse ”. (1017) 39min; 3wn~iflfih Ly“

[1.1; . :2 '14: Q

-__—---—---- (1935) An unusual Chimacre

Soc. 7%: 241-44.

'Yarnell. 55. H. (lfldfi;) Oak bfeélfilfld in T

~128-

rn . ;"

4176(48 Lg {WK-J

r.‘

ULZ; "1. C 133,51. E 1.1 i 1111 e l

ElJn btnnilbr

{TE-.19 £3111”: fHTHilzr inClLlC188 fillflllt 1F) genera \fi_th 11301.8 tlign

150 species of trgos, shrubs, and herbs. Peprensentstives

J.

are found for the most part thruout the temperate reg one of

(Ho

both hemispheres. A few soccies are limited to the trepics.

U.

y.these is herbaceous. 0f the remain four, two (Ulnus L.

H.

and Celtis L.) are u1f1o1en-;v common and mportsnt to be

considered of interest to the forester. The other two, Trrema

o.

Loureiro, a tropical genus found in southern Florida, and the

monotypic Planers Gmelin, restiicted to swamplsnds of t1e

southern coastal plmein are considered, at present at least,

of no commerccisl importance (Harlo' e Harrar, 1941).

The economic inhortsnce of toe Ultmseee primsrily depends

upon the valt1able timbers prod1ced by speecies of several dif—

ferent genera (especially UlWUs, Celtis, and HolOpteles Plan-

l nchon ) and the high ornamental value or many‘others (Pearson

% Rrown, 1932).

UlLUS L. (elm)

1

Ulmus includes about 18 srborescent species scattered

thruout e1stern North America, EurOpe, and Asia. In many

regions, as in parts of the eastern United States, the genus

is en importont constituent of the he rd11ood forests.

The genus is divided into 5 section, all of which are

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, also of importance in India.

asentsi 3y toe six native soecies of north America: (1)

' -"\~"~ h h - r‘ ‘ .- o ’- 1' F‘ V news ‘r ' ~- 0 f .l‘ f“ -

3163L8T04dlfius Dunn lnClU;cS g. a c11cgna L., (2) Cnaetoptelea

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(‘1 ‘- —= “P ,3 A 1' 1—1-‘— - r l . 7 " _ “pacn cciu :s e. crass11olla uttall, 11- (5) Fr)? uptelca

es ‘ f ,‘ ' ,_ ,r .,- rm 4— 4 ‘ r“ a

ocnnei er inalu c_ U se1u11na arrte t.——— -—________-_-____'.______. .,

‘ 1 r~ \ -.\ ‘A q ’ '7'? ‘fi "“ "‘ "‘ - '_ ' ' ‘I H 1‘ ' 'Tne 1 /-” lelUQGS Sufie or fine most uSeful as; “811— rd.

t3? six s,ecies nitive to 1 : UchEl btates, t. awericsna, U.

to0“2si, 311 U. tslv” "re of ‘rvetest iaportllce as forest

JG ‘tim-

bers of all the species since they are all usei for essenti-

llly the same ourcoses whic1 incluie slack cooperaee, boxes,

ness of elm wood is adagted (Brown & Pansnin, 1940).

(u

tne elms are UNSdeaS"[1"

the Unitei Statis, the List 1 oortant are Lse nycn elm (g.

(glabra Hudson) aifi Enclish elm (g. orocera Salisbury) ofz’

1‘

Europe and their numerous varieties. The Juinesc elm (g.

cs 11‘ ~1

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has been wijely a“ sue essfcllv plartei in the plains re-

Cion of the UD“CC‘ Stites fur 1113 breaks an1 ornamental

puraoses.

AETIF"?1E 3:1;1TICN 9?C1 MILD 5103K A.b TL?T“CL-§D 3>C5338

Selection ani breeding stuiiis in 1:3 genus Ulmus have

been greatly stimulated in r ce1t wears, especially in nurses,

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I

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8111URS 11_t-ir uLc b.i33158 111 re an - to stwnaeotln.111t1r‘t0\ ~ ll

‘1‘ _ >gV ... ‘ — _‘ . _o .-.. ‘ " . ‘ fl 0 I“ '_ -\ -. r" “ 1 p .’ \ . ’ x . I I

tie oisesse. .o SivaC 5 60108 u: AmEIlCd, surOge or seia

:as been founi to be inxune but some, as ecially tne Shinest

or Siberian 811 (Q. Jumila L.), are usually very res stan .

Kany varieties or ferns of some species hhicb are sonsiiersd

on tn: uncle to be extremely susceptible have, however, been

proven to be resistant.

host promising 2f tne highly resis ant clones isolated

to date is the Dutch variety designatei as no. 24" or

”Christine Buisman", a variety of Ulmus foliaceae Gilibert

(Buisman, 1956). Cf soecial interest is tne fact that this

species, on the basis of nany other tests by various workers,

Was consiiered one of the ”very susceptible" species. TheOI

”Christine Buisma " variety gas been tested by several work-

wjg; sent, 1938—a; and-./

l—J

ers (Goidanicn & Azzaroli, 1938,

belch & Collins, 1940) ani Has been recom.enied for culti-

vation.

1

Presum (I‘1 natural hybriis in tie genus are few. Johnson

(1939) listed three. Of these, the most important is g.

elabra Huds. x g. carpinifolia Gladitscn = X E. Hollaqdica

kill. Under the latter name, Rehder (1940) included as var-

q

ieties a number of long-describe1 plants of variable taxonomic

rank, such as ;ne "Dutch elm", "Huntingion elm”, "selgian

elm", and others. Taxonomically, in View of tie present dearth

According to Bender (1940) this is a synonym for the smooth-

leavei elm (g. carginifolia Gleditsch).

-132-

of cyt genetic eviience, such a metnsd of grouging is perhaps

justified. rr«SuraLly such grouing metgads are n:t designed

to indicate tit: fiiality that the classification is correct

or to infer that each cf the varieties re;res=nt members of

F1 generations. Sax (1933) concluied twat the numerous var-

ieties of X g. hollaniica are presumeoly sereg tee of sub-

sequent ge:ier ations. Two of the varieti 8(E. h. ritteursii

(Kirchn.) ehi.sni g. h. sneerba (Aorr.) send.) tere studied

cytulqtljaiiy and revealed Wnly sligat meiotic irregularity

‘ ‘ ‘.- — A “-~ \

other issuers Of$.)4

an; hiya ,ol‘en fertility. Sax also stuiie

P“)

,0?“A.the genus an: concluded tnat a number 0 tiIiCial sgeCies

hybrids could, in all erobabilit , be easily made.L

Henry (1910) studied progeny of flflat he believed to

represent the F generation Hf the g. elagr x cargirifolia

2

cross, i.e., offspring of two g. h. veeeta (Loud. )_

_1

(led.

individuals. He reportedly observed three characters

(oooosite vs. altern11+e leaves, leaf size, anzi lensth of

petiole) each of which segregated on a 3 : 1 basis.

Artificial hy.riiizeton in the genus was first reported

by Klotzsch (1854), the ”first tree hybridizer”, who succes-

ffusal. The parentage of(D

sfully crossed g. camoestris x g.

ctional(I)

ate an inters(3

this cross, if correct, wou d iriii

cross: 'eloceri“s x sleinaroowrous.

Sax (1933 resorted a possible cro s involving g.

americana L. x Q. la vie Ball. The forner is tetraploid

(4n = 56) and the latter diploid (2n = 23). The ofspring

would ha e pre wesc‘ly been trioloid. A cytolosical report

Probably Ho slztra L. x g. laevis Pall., see Rehder, 1940.

-133-

on the of rin has not come to tne sutior's attention.

a lieite d amount of intra- an interspecific hybrid-

k‘:

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F C p.‘

O

J

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FJ.

:3 h (D

(B

i H O D ...—b

C H

O

"a“ 1')» r 1::-

Cu sarr-b~ization tork has 0 (D

a disease resist1mt form of elm. Uoorznbos (1933) reported

W D U)

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(.1 u

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r—

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DimCl‘V: x .al ic1iana, T. laevis x 10113“18a, d“l U. Jumila

Smith & Nichols (1341) reported tne probably successful

crosses: Kg. hollandica hill. x g. iaeonica (Rehd.) Sara.

aid g. Hilsoniana Schneid. x g. *BDORiCa (Rehd.) Barg. A

resort of tie eomuarisen 7f vigor of the eresunei hybrids

with their carents nae not yet been made. These torhers,

hutever, stated that hybrids of the Mtter cross at least

apoeared more vigorous and more realistant to iLS€Ct attack

than the isle parent (g. auonica).

interAtterois {o obtai involving Ulmus3

W ’_.;0

O O "3’

O U1

U)

(U

(D

azi Jeltis have not been restrted. Both _enera have the

same basic chromosome number (14) an; if comsatible should

yield interesting hybrids.

Flowering habits and I§Vfl“’izlng1technique

The exceedingly small aid delicate perfect floners of

Ulmus are borne in dense fascicles or racemes which maies

emasculation by forceps an extremely hazardous undertaaing.

(-‘I

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l V O Q I

Aut orities not cited.

-134-

x 5,. 1. “- ° . o A..- A RA , .- , -. _... .1. n ._ , 15 - ‘

is so tar cu in sun; s —cies at least tnat ejasculatiin is

not necessary. Last (1940) also noted marked protogyny and

V. A‘- ‘ p ‘r r ‘\ v ‘

b duff." . Y-:2’S 3.0. ut- :l’ ole/l u 11“. sci-o .‘3- OUL. l".

l. . v " ~ ‘ T p - ... _-‘ , j ._ _ . .‘ f_' v I' . ..‘ .‘ -‘ If

tiis genus. 11 mass emaSJULaLl n 91 eln -louers coul i be

w. . ‘\”‘\»'q ‘ ' ‘- . ‘—~ “01" . r r ‘ " . .-

aCCOQQlIS'ci Mituout injury to tne resale organs it would

e oossicle to avoid undue worry for the hgbridizer (i.e.,C,

.L

worry over the existence or not of effective protegyny in

the sarticular female parent beinc used) and incidentally

give a more reliable ani imtediate indicatifn of results.

The hybridizing techniques used :y Loorenoos (1238)

H.

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unfortuzwate My ha e not been a ailable to the author.

Smitr & Nichols (1941) resorted successful use of the

"greenhouse techniiue” in their elm nybriliz7ation more. This

method, first recorted for Salix by Yanchevsky (1904) and

for Pooulus by nettstein (1933) employs use of flamer-bearing

I

cut twigs on which Iruits are successfully matured in the

1

escrioe Q.»

by5.:42 . r .

greenhouse. The metgo; Ul 3ollen storage a

Heirwur""r (194C) i.e., storage of ocular flower bud-bearin5

dormant twigs in refrigeration, maytwesumably be used suc-

K)

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on techni;ue, dalix section c: this re sort

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for 1

q

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4

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11

re lists by var-x

.- -

m

chromssm

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V

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.

(3

LVWm " r.- .

ILA [-1] L) O n' L

ious torLers (hrsuse, 1S50; Sax, K., 1333; GdiS°T, 1933;

T A . , .7 {\7-v - , "‘. , - “" ’2 ‘3 n - 5" "7n : .",_‘lellvelu, 1),), 1955; 01k, H. d., lJJU; 281 nauie, 1259)

T‘L f‘ 1ffil"'(" . ‘$ ‘v"' A -'

LL18 5.5.013 1 LA-ACL:I- .Ln

r. 5 .N

neterUQIOiog

CL:

*1

'(z

1,..1

C-

H CJ

Cf the 15 soeoies tgus fer counted al; are

(2n = 28) but one, he Ameriovn e1: (3. a erieina L.), WhiCd

'J.

U)

('4‘

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(L

H S'3

k)

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0 }Jo

Li,

A L-

5

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U1

°A~ «0 .»-, 3 , .' ',..,... ,. —.- . Z:».‘ 12..., ',.. -_— Na

tlun u; seco aary 3a1f14» concluiea tue s3e31~s has grop-

(a:

:1

m

. .V". 47 J .1 . A , .. '1‘ 7““. 'p ‘1 IN 7

tion, is ettrioi.e; .o litites Zulab - ircoqenog.

Sex (1335) reported only sliggt irregularity during

meiosis in p.m.c's. tf varieties 'itteu:sii (Kircnu.) Bend.

" —" 1_. ‘, : -'~ 1‘ ~‘ ...\ n‘ ‘ 1 . “ . l‘ , .‘ -.

uo-rba (Lorr.) semi. ml A T. pollennios e111. uni con-

OY‘» C

’AL.’ v

_ _

‘v : -.,. .L‘- A- v. ..A ~ -. i . J. .. . .. .- i. - w - x-c‘ ‘ ‘ r .T' i- A

CLUlta -Jey msy zeoresent later SEfIZtALCS U1 nyoriis oe-

‘v Q

olia aleditseh. Leli-

H 9 C.)

£3

(L

VBli (1933) Uts=rved regula- :eiosis in tge rari t; oe

J- “ \'\ a“ -\ - - AA ‘7 " ‘-\ - ,3. " ~

HOD S‘p;urt \.: .ucurt a- “yell“ UL

v-fi.‘ -. \ . ‘ ‘ . o o -- I. [3 o A _ A“ ‘ ‘37 K 3‘

lee aogarent Sterlllty o1 LAlS fozm has Conclaiei sy th.s

;.

SJ

E E.

Q

author ( 955) to be loe .0 its extregely marxei

rat*er t;an infertility of its 9; etes.-A‘J

nalher (1952) resortei ine relucej number of 15 for g.

oumila ani U. fulva an; 28—30 for g. a erioeua based on

stufiies of EicrosoorOVenesis. The sine “or; r (1958) re—

oorfied 15 bivalen s ajoesriog at meiosis in to: ovule of

9 g. fulva.

‘ The variet" Q. g. oen€u1a Aitufl has also sol iej b‘ Sex

(1933) oil f oil to Le tetraploii.

' Truv 7~ 'I’T‘Z'J’f 3" air“J1--.- .11)-"-qu 'JI...-A-L)

(77141; C: )

Crown, 3. P. ' Pefisbin, A. J. (1940) COflfiercial timbers

of the (wired :Istes. st Ed., AKI+ 554 oi., 312 fiQs

LPT‘Ww-'il: Rook Co., Inc., flew for 9 London.

uismin C. (1”36) Le resistente lea Er. 24. (The resis-

tant elm LC. 24.) Tijdsobr. fied. Heiflow:stsoh. 46:

:3—76, (Cited frow Elsth Treed. Abstr. 01. V , 936 )

r3 -= n.'ixzk, l. '3. (Laidtrteci) lflfik:eiz oi‘ s;iitle:ti<: Qrwy t?‘ sin)-

stsnees on tfie rooti{Q of cuttiIQs of Wood" orruNIouql

plants (“IINLllbefi).

-------—----——— E K1311EVOP,L. C. lffig) The efiget of

synthetic Qrowtb :;u:stniees on the rootin' wno subseoue-

nt igvxrth of cmqv“wgnizfil plants. CNTlO 5X55 .flra. ‘Fimorfl‘u-

1y 5H1”. 24: No. 199.301unhus, Ohio.

DeerenMos, S. 1. A. KemiserSfNoev01 met iegen te ‘s-Creven-

hege. (GrossinQ QITCIl‘.110ItS Ni‘h elws fit the Hague.)

Tijéschr. Pl"iekt. 44' lol-o4. 1935. (Cited from

Plant Treed. Abstr. WI1 IA, 193?.

Gaiser, L. C. (1955) A list of ehfiowOSOme nnnbevs in An-

Qiosperms. 17. 5i} ier. lenet. 10: 135-258.

Goidnnien, }. (1956) (i‘he -GICiViORP of "U. pvwila" L. in

greetieel F Hricolttne 0rd its Pesfstenee to Greyrin_m )

13011. ~+s" lfit }%~’ LTW“) 1C: (I1 3.) 11% ~2CfifTEET7CEE1

from Plant Rreed. vol 7, 1237 .)

-----—------- (193s) (Notes on r'searehes 1L the selec-

tion of elms resistant to elm disease.) Itsl. A“r e.

75: 69-74. Cfter" from Plant Tweed. A s'r. ”cl. e, 1938

——---——-----—- a £2 eroli, l. (lQSU) Pelszioqe sulle esye

lens di selezioe di olni res tenti slls Qrsfiosi e di

inoedlsziohi nrtfliii inli di "Craphium ulmi" es eQ11-e nel

1957. (deport on ihe ox_reriwents of selection oi elms

resitfimwnt tun elniciiseese=rw C of‘rirtiiiwiwl incwyllsticwu3

01- r}. ”3“."1 izwne in 1337.) 01].. fist???” Pot, V33, Young

1L Tn.s.):149—7e. ‘

--—--—-------—-—----—-------- (l 3”) (”eport on experi-

outs on selectinQ eles resistan to

srtili Cisl inoculations with G. Ulmi—--—.

Plant Freed. Abstr. Vol. 10, 1945.)

1;?) Plow, , V; 0 iv. o ._ 113.71 'C l1 , I}. o L) 0 (19/1'. 1)

05y. 2nd Ed., X\i-+F4 pp., 234 fiQs.

Co., Inc. Lew York & LOPGOH.

.imburQwe , C. (1?4 ) Report on Peplsrl

CrorestryCironie1e 16: 149—60.

(‘

seedlin«;s

(lmn:dcn2)

tnel

O i' .9 1’1

elm

SOC.

ru)te (n1

oy means

(191C) On :‘10.

Jou r . Linn .

(1027) A.

iloxens

ITE‘nrjf’ PIA-0

sults.

Hill, A. G.

g. 71‘)

11‘) 4'”

x'v Kl;

Sly-U371— as

ilQri/Wilturwe,'lrfl11. 14: rye.

KlotZscon, J. E. (lcb4) Ueber die latte

zen- strwoe one iischlinQe. Her.

Pal-Led. 5. ss . lfierl in 1:. 54: 53-5- .,:62? .

-138—

fextbook of de

)’?10t.

ed for

nirstor

1.701)} lglld l o

(T:el disease and on

in 192’“. ) (Cited from

ndrol-

Lofirsw~flill 500k

‘Vbl‘1.61

in; mendeliqn re—

59; BOO-SOC.

sculeting

from.

em 9

Cor Pfen-vrehour" -

Meme.

1."

LL 0

Krause, O. (lQEO) LtoloQische Ltndien bei den Urticsles.

Ber. dent. botsn. Ces 4’

“e

. 0: ~11.

Krijt}(, Ii. (”771) (neport cul'tiie tiviti iVHJ the elm

61385188 curm‘ittwee cs W‘zied.cmxt at. t}ex ene:1i.rrs Ernie/'r‘niry

in 1937.) Tijdscnr. PlLiekt. 44: lFC—CO.

-- -------- - (1793’?) (iegort on tz'e activities of tie elm

diseose conmittee carried on? 't the rcmetics L«oorxiorv

19L8.) Tijdseir Plgiekt. 45: jS-VO.

'Lelivgld, J. f. (14313) TCytcloQicsl stuCHfins in the venue

Ijlmns 1;. I. The ingjposcd Tnfiggid rwflnxre CH“ tre (x) vuon

Eaten elm. Genetics (Etc VSQUC) 1F: 4h5—15s. (Cit -d

Iron Plan: trees. Abst‘. TVol. 5, 1355).

-------- ------- (1‘35) C tOIOQlCfll studies in the genus

Ulmus L. II. The entryosqc and seed devolOpment in the

COquH IDUtch elm. Rec. TrsV. Bot. Leerlsnd. 32: 545-

573. (Cited from Plsnt Freed. Abstr. Vol. 6, 1936.)

19y, Curtis (lCS4) outbreaks of tie Dutch elm T'_csse 1r

the Uniied :tetes U.C.D.A. Cir. 382.

----------~ F 2°“Vrtt, E. B. (15"1) th L to: eln di . se.

U.S.I.A. Ci". l"C.

Peersor, R. S. A Hronm, A. P]. (1932) Corrercial tivbers

of InCis. Calcutta, Cove TFWL of Ineie Central IAiclicst-

ion Brenct.

Render, A. (19/10) IT.LFT._T:U.91._ of“ clii‘migd trees 916 ghr’u‘os,

2nd Ad., 996 pp. tscLillsn Co., Aew work.

Seste, D. A. (1955) A field aspiiatoi for p.33 culstinQ

sweet clover flowers. J. Aner. Soc. Agron. 27: 774-

75.

Sex, M. J. (1038) The relation between stomsts counts

and chromosome number. Tour. .Arn. Aro. 19: 437-441.

83? Karl (1%?35) Cowomosome rumoers in Ulmns and related

genera. Jour. Arn. Aro. 14: 8/— 84] ‘_’“"

gchggrz, E. ( 737) Ist die Ml tterulme res

Ulmus lsevi s reis tent?). kite. dtscr. dendrol. Ges.

49: 185-86.

Snith, E. C. A Nichols, C. Jr. (1941) Soecies hybrids in

forest rees. Jour. Arn. Arb. 22: u4Q—454.

Stephens, J. C. & Ouinby, J. R. (1955) Bulk elasculetionC (

Of 801231113"! llOWeP. . TOTlI‘. Amn SO . 5" 51mm. (35: 2.5.73-54,

Cuneson, C. A. (1937) rquculfltion of hnnit by cnilling.

J. Inner. iflDC. ingror. r39: 2347-4<T.

Tuheuf, C. F. von (1935) .crae"sn, der Erfors chun.Q den

sog. Ulmenkraankheit in Barons. (The deveIOpment of re-

search on tne so— called elm discese in EurOpe.) Z.

Pflflrankh. 45: 49-78, 161-89,

alker, C. I. (1953) Chromosome numbers in UImus. Science

(n.s.) 75 (1E34): CV.

-—---—----—- (l 58) ~TCPOSWOIO‘:nesis and embryo develop-

ment in Ulnus Tlvs. Bot. Gaz. 99; 592—95,

l

‘cu

Went, J. C. (1958) C AnilsLion of

Aelch, L. A. A Collins, D. L. (1940) Luten_elm disease

end its control. Cornell Ext . Eu.ll. A0. 437: Pp. 19.

th investigations on

14%) mjsceotioiljjiy o1 diiien‘wlt elms t1)(yuu9tostowe11

nlni Biismen in tie Netherlands. Pfiwto) tn. Z. 11:

Iel-BOI . (Cited fr Pl. Breed. AASLr. Vol. 9, 19:1.)

'00

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1_7()YI

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LCJIQC.

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. \ . _‘ _ _ ’

VI.-

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stijations on elm

13011019 en

1936.)

'F.'I

(1111‘11’11’. 4- ”u

(dited1?. from Plant 3096.

EJ";66) 1310 IICrWIzuI ,snwatluJie

T95+C :Cde -Ci P0pI1n_s lei”-

M1rtun; a: 597-026.

Cecilinrities Of the Servin-

n i of some 8000568 of ”illow.

afiest Just. 11: 209- 274

10L of ‘reon and shzuus.

1FI'7. {3 pl).

?”"CI-”ED

., '1 1’

J. 9361:. 530: .7341.

Quart. Jouv. For.

29' 102- ”.

Funk, 3. (1351) "oer Cie ?’”‘9tifln CC? chbkorren Lweier

vanaschierten UlwI;n. hitt. lent. Lcnflvol. @939911. 42:

525-528.

I‘fr'ijtbe, N. 3: 911.2, J. C. (12759) (II‘OC1JT1—a1'.i()1’ls of elm

Enfl391ds, CP“"ried curt in 13%?3.) TWduisChr. IfLI”€1ft.

’5: 1-74.

Navasbin, S. (18TC) ”he? d9: Terh51ten des POL1ensch19u-

01%28 Ewefi CI>P IVH'e. PUJ 1. Jlflfic. I If. F*t. IVsteIwfljvlfg 8:

545-n58.

"ent, 11. C. (1”"') Vermilsg 'Inl Jo Cumierrcmfl:1n¥p11 OVCI‘Cie

:_e:‘=s",:1".7i€$ 12,6 VGI'I‘1_I:J'“17 Op 119,13 EV’IUJCOJ'YJLE'C‘lOJJ'jJBF/‘h Lt"I1“IC\:I9-

Cerium ” ilJie Covmelin dOhOlLCL” te fianrfl, (~019nvde

1977. (RCport on the investi;9tions on the Ele 618 539

C5PTin out fit the {11119 Comwelin tcholten P‘;o095tho-

1 109 L5bor5tory 9t R99rn, durim; 1937.) Tijdschr.

6NC

;:40

930

44:

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I\ (A .1 1- e r 3. . 11~ I),

m1 , r '1 n :5 ‘ h 0 «,- 7 _ 7 ‘ ‘_ _ w _ P ‘I\ __O ‘ 0

J33 lglv -, 111 neluICs Cut i.o :2:13: 'J;LFYU“13

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A. “can (sarlom & flilrsr,

r-"Q'1 , ‘-— -\ r‘ 1: Y'. .— ~r\ ,: y‘ 4“ In ,\ NH.

198 Celt-anoun euJ must valuJC(II

genus is fpe sumar figple (4. sacchsrum) which, togetner with

;~ ni¢rum, is tIe ,rincinle source Ml "harm Leple” woud 951

maple sues? of th= America: trade. Hood of hue Carl :9; es

is fevoreu for a multitu.e u; usss, lfllvf 95035 LdiCh are

l . . , . . l_ .Cne spe01es, noaever, 15 the mountalns Cf Java.

‘H ‘ . v! n fl T _o a If _C

LchHOCQ i'bLJ1L3t

W . r. _. “ V‘ o r‘ _

__'.,1 -141" .11.) - -i. V'Tj';

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wooeents'e, HLVCJ

(ETCMI? i réifshiri,

The "soft 1

utilize 1 Car-Ce L3,"

except Lnere stre

reason soft “Hole

Cregon ra-

veneer a: l m:—

Vslue :u. Lu wn~

maples.

"A .1- ., 4. fl

,L.L.ISL, (J- uL;C

Hum er of exotic

as ornatentcls,

sycauore us:le (

A ‘ ~ ‘ - f T "

cclucstre L.) J

D - ' -I

0; exotics in tn

'$

future Cree

1- 0 ~__ ”A

stocts in tee t—

ies are ;n ,ro V

Station (:ohrein

:C

.1 .

.- LL , ul ,

“ : .V 4. -. D . -._ .. . , 41"] ,-

; v-:eu LCJ“JJLu1€, JULUO

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12p, Q‘ Nd ~rous otgc

"‘1: l \ ‘2 , . . - __-~ ,.

1:40; £83150 6 glam“,

.. ll - x “ - _.- A

~_L‘ .1 (Ti-PIWrI‘Y'I.‘11nt);

v L.) -. . Ugr.\/ .JLL A ‘. .1._A v\ k -4

"I 4' ‘ r 3 _ ‘1 w y- ‘) CI .2 C.‘ 1‘.U L}...ka UMALL /L(_‘ :It uyu g‘.

[-4- ~r‘ r) "1 7 ‘ I“ a, 7"" r~ h r- /\ -~

.__ L... .:.. - .. ...;L .c ‘ :. all C t.

I .u‘ a A - ‘fi ’_ _’ 4") .

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N . «xv-1'11 ... 1 ° F.

ii 0 (it ‘4. ~13 C' [.1 . ’Lv ] l. 1.1:)

‘ ‘ 1 '13 ~° c‘ 9 - "1 + I" a WI [l4‘u..1 .LS llue.e .1--1 U-Av L;.

g2) 18 O? .ZUB

(74.1

LLAVNative of

tritution.

1"; G.A.-

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LJ

the

-. 7,) K,“

"' ' ‘1“.ACCOI’

others,12')

~142-

to bcnreiner,

K J

C. ' ,

1;

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S'ioneole

14.5

me

boxes, crates,

oil" ar+’cles

).

{lgeruxfi 'Ire

asd fflfiliE

I \O

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1‘3

1.4

O

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lf-

or floor.(1)

rimarily for

snufacture of furni-

1~— r— —‘ , ..

C 01.1.-LC 1 C .l- E.

f“t¢e CCMR Crx:st, a..1 Cesat Laiowrl of 'tse

ative ,e Cars of is; enus, a3 well as a

;;ci;s 97? varieties are tiiely plants;

,Coicllv [orwav La‘le(A. olctanoiies L.),. . _ l.

pseuioglatahuv L.), beige maple (é.

ficngr:uther£3. Sucrlei no CILMJS o”d_t=ct‘CXI

country shoslfi fireatly fascilitcte

r.; in ixie gxyius.

l

ZIJL iglfJTI H F-C- u: ~ cICJE

le regortei evijence of selection from w’l’

3 other than tse indication that such stuc-

northeastern Forest fixperiment

studies

7 ‘r.‘ C V

A v La ... .

\ ~ A

“.»“’\":: . 1f-1a'x . ‘ .'-§r - ~ »4\*‘ v-xw" -7 -.-. .. a)»- . A.

1:- to isolate suierior loros oi so-.r $1218 is: su ar

grofiuetion have bees initiated.

lO tattra tuftr14s in the .eous n.12n gave be;n recognized

by taxonfisists, out some of trege, apgarently, have shown

special value for forestrv use.

CL, f'l'3§,l._i.i.;:; g‘.?.:...'31:3 ALL. pal; If- ..N

Only one artificial hybrifl has been reorted in tle

genus (Freeman, 1541). This cross, Aoer saccharioum L. x

Joer rutrum L., yieliei 41 byerifls ngCh shonei consider-

able variation in leaf form, habit of growth, and hybrid

vigor. All of the hy.riis s:osei markei heterosis as

comoared with the male parent (i. rytrum) but none equalei

tLe raoii grouth of the silver ragle (E. sacs...r1UJEIM)

fiourteen of fine hytriis flowered iq toeir 6th year, 10

H)

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feeuliarities in tuC flowe;'1rH Habit of tee maples ago

the lack of hybriiizing technigue is ,rcbatly responsiele in

lar 6 measure for tne aos=noe oi artificial hybrids.

Sohreiner (le3-a, 1933-o) has re'or.ed stulies of Ith-

eriqg natit is t e Cu-”v i . rej Ha.-e tutor is of sotoial

maple revealed a tyoe of elea -eut iiohoEamy. an tvpes of

flowers are born by unis tree: (1) Jerfect floners anj (C)

imperf=ct flowers. She .2err1:ct flox. ers oere ooserv=a to lif-

fer fro: most bisexual floners in that only tJG male or fe-

Individual .rees mere also note; so be eitner :arkedly pro-

tandrous pr frotogynous. In the case of grota dry, for in-

stance, some gerfect flouers nature; t eir anthers coinci-

dent titn the imoerfect male flowers. lhese oerfect flowers

did not function as feaales later, houever, but other per-

fect flowers 31th functional oiatils did. The periods of

n.l\

\-

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I1..)

polien shelning an; stigma receptivity in a s e tree were

not observei to overlap ani thus cross-pollination resulted.

rr'v‘

1ne flowering habit of the re; maple (é. ruorum) was

reported by Schreiner to oe more comolex. ’hree tyges of

treLs were ob s erved: (1) all flowers functionally male

only; (2) all flowers functionally fe dale only; aid (5)

flowers functimvnally ferale or male.

U3

ome trees in the latter csteHory snowed predominately

ma-e flowers in the upoer part of the croun; an approximately

... w

lequal proportion of male and Iena e in the midile portion of

.1

the crown; an: predominately ferale flowers in the lower part

1 a

of the crown. host of the Ilooers on all types o trees hereto.

oerfect, but the non-functional parrts were

silmlaly smaller or aborted. In s;me instances, however, func-

J."

tional .ale leters vere observed Which had no evidence 01

1 Trees (1) anl (2) were tnus furct;onal ly dioecious; (3)

“as fvnctitnally uoroecious.

-l44-

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.11.; 1’_l:,o;.ll (1:3) 3.." _, .-.-:4}. (if-4Q) tjtlirjlul'} tLlL [8.11.1.8

a n, 4' ’ ~r- c (3 .2] f 5' "3v * ’1 1 t: r: .-‘ P.“ ~ ’~ -‘ "- "~t' -- 7-3 .‘ 1v ‘_~ ' w .-—- 4’- ]. I)": r: l.“. -1 v. LO wk. .1., .. -- L- 1...._'\: LAC. Lat]. cab -.-T1uh.l. ..Jbllj. L11; ‘ D’JKJ“L/ ,

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— ~. ’ 7 ' «J- " 3. -2 , J- ‘ J. - — .‘1 “ fl. n ‘ r‘ ’1 -\ - ‘ r‘ 3 *-

IL] br‘] , 1“ -11 J" ‘JC ‘1 + S L) k, L; _ S , :- .. .. A; t, , 1):) yr L’U --|.L‘7 U24; i.-

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.e--ers L1 1.: lenus aler are use; ly .ronpel Wltfl the

ill icult” g"a11s to root from untreatci stew or root out-

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in: (55.35 is lCB i vs) was ob aiiel with greenwood cuttings,

. 2 . .-- .made frno young trees tee miqnle of June. The cuttinfs,

4 inches in length w’th all out up er leaves removefi, were

maersei to a teeth of 1 inch in a Later solution of indole-

tutyri acii (concentration: 50 mn./liter) for 3 hours.L— I

To ‘° . -... ---. .Lg ... y. , .. -, , .~ ..-s - .—‘v.,~ .J- 1..

rollOWing triatmtflt L.€ cuttln_s were glanoei to aoouo LWO-

thiris tneir len tn in a oeat-SAn' mixture in an outdoor

} See also SHOL (ljbl).

8 anct are not s.ecifie* aLt10u"s, aCCor in t Thimann &

Lolisle (1939), this is most critical or in~ “actors which

influerce rortine resoonse.

ior 24 hours rooted well (65; in lC8 days). Suttin

after the mi.ille of July failed to root.

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tUbel Jr O=C Mari d—d SuftMUUi cuttih s, ugly one formed

r004 C‘ . t" : ‘7 T. " o (‘ ' ’3 - r s '3 C.‘ ‘04— , . , a 1 r1 ' .711, cu 1n 05 Jena 8. .LLilo URL y.;._b {A VOL LLVOOJ. htti'1; Vin/112,11

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cl lei tlat thh a lonrer {Ioniht season the cuttih

have rooted. Afahsie v' s cutti _s here is} in tie latter

I 1 ~ 1 fi‘ 4- f' \ ... 4.1.-‘,—\

r account in cart Lei the failure.

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Afaniiev gave no ihiicatien of the ate of the trees from

'.- .1 , ‘ .- h, . - 1 .. ,.. - -. ,1 p-13 - ‘ , a

HMLCH hi: c Lbiflfib were haue uni it therefore agoears pro-

aole that rooting failure is this case uight be traceaole to

Show (1939-b) founi optimum auxin treatment for scer

rubrug, pasei on 50 ramidmly selec tei clones, to be 6 hours

in a 200 rag ./liter solution of irloleoutvri acid. Untreated

cuttings are saii to seldom root more that 102. Of greatest

ion(1+

significance tere the results regortei of an investiga

into the variation in the rooting re-oonse of seveeral dif-

ferent clones :F rei raole, :ll of which were auxin-treatedw

in ‘he same manner, i.e., 6 hours in a 200mg./liter concentra-

tion of indolebutyric acid. Variation in rooting ability

~ 4“ a A l:- fr v": r} .-. ~ g“ fl _“ .4 —-. A r" 11' . .

Jiflt; irom l{.§ to >7.§s along the seVeral clones. upon the

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‘ J— , A ‘\ ‘—-, f\ — : ~, ‘A .3 ‘,-. 1 v 2 x «:x -.. ~- -‘ "q ‘ n 3 . f‘ - 1"

fOYEEt art: ereeaer it urbwaplb TereSugub an: lirot kaif-

‘v ' 4- r w r: "\ ~‘ " ¢ '2 " ‘ " ," " ‘ V, ‘ (x a 'V ' ~ 2 .1 -~ .

1 ental EVlaenCc recoricl in suugurt uf the assumei renetic

founia ion :f the "rooting ability chaLacter amonn furest

;ians1ev (1339) also reuor el auxin treatment studies

1“ " 0 ""1 “V ‘2 :v \ 'T‘ ‘ " ‘ " ’ -. vfi r ‘ "‘ 1 4' \ \fl'l . ‘

of rea v&y_: CUtL n_s. lflc test root .2 results (6Le in~-

iays) were obtained Lith greentooi cattings tale in early

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$3 IJuly anm treatei for 24 hours in a 2C mg./lite;

t_on .f inQOLehutTIic acid.

Thimann & Delisle (1939) investigated routing ability

of seer glataioiies anl the results ostainea lidicate the

e (J' the tfljart from.wtfixx1 cut-

tin s are mane in jeterminatian of rooting aercentage. gn-

treatij cuttings from seealic s l/2 year old gave 67% root-

ing, 2 vear ola plants gave 4Cfi, plants 3-5 years old yave

”“1, 4., .Afitfl: .“

;1ares root—u only oc-

o ' y 7’

cassaonally (less t‘an ls).

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Although untreatej cuttin s of 1/2 year old - .3

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3 lrooted in 67% of toe caI

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are-.2 L“ —. . L-, f ,, f1, , 1 4.-.,“ ,

Qractical f«r fUtEstr; use. Jutt 1 a from trees 2-5 x'-

old, ~uwever, saowej 44% rooting “gen treated with a 100 mg./

-147-

liter co centrat on of iniole—Bacetic acid. Cther invest-

1'} :

‘

‘ . 1* ,‘N w; J- : .'~ J. . - .,. ‘ "\ n! J- ‘a .— f -. ‘1 - - -~ ‘4“,

.11Cateu that cuttin s Iron the basal aarts of the}-Jo

ifiations

tree were best.

Chadwick (uniated ani uuoublishej manuscript) obtained

2Cfi rootin: in are 42 days of cuttings of Acer galmatum

Thunb. treate tith a 5 mg./170 cc. concentration of indole-

butyrin ac1d. Control cuttin; of this soecies groducei noU)

‘

roots. Acer camgestre L. cuttings all lied without root

proiuction when oiven tie same treatment, as well as sev-

eral variations of it.

”ffCuQGY(J

Chromosome number

From resorts of cytological stu

by various norkers (Darling, 1909, l§l2, 1323;p

U)

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an interpretation of\\

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some 1 wer number forC,

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mate basic number of l” or o

the genus. ‘bservation of seconlary oairiu: has also re-

coried by Foster (lggi) in 5. nepunio var. interius (orit.)

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011 (2n — 20, as a. saccharum bwéytOlOgists, 14 are dizl

Mar 3;. .hi “. olatonoiies L.); 4 ar= tetraploid (4n = 52,W

'- .’. .‘ »‘ vx - ‘ /\ . 1 .' ‘ D I" Q A . -~ ,‘ . .... ‘ - . ‘ ," ,‘ 1 r.“ “‘ -| “I . ' .1 7“ _‘

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gequs at- both rearrteo "QC regula occurrenc o

chromosoie icu l1.¥ in regions of the root_tjps, In the

case of the triploid seedling notei abore, Jeurman obser-

73 number.<1

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Chromosomes of the octaoloii, é. 5ubram, have been

studied by sev ral workers (Darlin 191“; Lottier, 1914;

Iaylor, 1920; and roster, 1933). The reduced number of 40

‘ _ ‘ A“ V ..f“ ‘ .‘ f "‘w t. “.’\r. c ‘_ l‘ 2' :0

Was re}orten oy Darllflf; 50 by rottiel, o, a 90, all oo-

75 by T aylor; aul 52 by Foster. 13ylor' s j 50 agioroxi-

. V - ..C‘ ’:’ .. I 4* "w '3 ’- 4' ‘rfi ." "~ 3,“ o“. 7

mates roster 8 count 01 p2 agl onus 1avoxs the oroeaole

Tooter concurred Lith Taylor's suggesti n tgat other

forms of the red resis, having jifferent chromosome num-

S-b) Work with the

(T

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he 10 of the 14 secti ns ani about 1/6 of theN ‘,’- J‘ .

anotn svecies.

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Vgrl&tlbd o: to: rowtlnt 'cility oneracter in tul

may conceivaolv be corollatei tits coroxosomr number.

“‘4. . . 7‘ " ,V. ' .. , 4- \

Lg er citolotici motes

Liloto (1529), Leurman (1333), all Foster (1935) cote

!

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{I)

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1‘] 13 _L‘- L. .L UI :11 L. Sa,--‘JC l'.‘ E) {ELK/1.1483. c 34175.] 4.11 A.

...,

-un seq other polyoloiis, no univelent or multivelentH c (‘1

w p

SD 0 (U‘

(J,.‘

O iations were observed. Foster coxcluiei teat bfliS w;

, \.~ ,5. i. -. nusm. 4 .v. v :1.” '~ 7- w‘probably uue to tUe low Cflldcmd freooescy uni)“ is grooably

" r‘ ‘ ‘ 1" 'fi " ~ r- "L ‘n . v u I" ‘ 7" . x -‘ . \ .p \ '3“. - ‘.

has OoberEQ by tLis tors—r afl; also 1...eurulen.

Foster (1933) reoortei collen sterility counts for 53

soecies an: varieties of ragles. LOSt o them (45) showed

more tgai 83% goo: pollen and the otuer 8 veriei from 100%

sterility (é. teggentosun Laxim.) to 257 sterility.

-150-

1'1'“7_" 711’ 77317“ ’1 ”11 Y

JJIR'] 'lz'f-Q YJI-.L'.‘on J_J

ffcnsicv, Iichcl (1939) Effect of iuflol~211ygiv s-id on

rootjflg3 of ijywrvo‘o v7'133133 Ol'Pm)W3 decid ants fores+'

trees. Jouv. For. 37: 57-41.

Brown, 1i. P. F FHWislin, Q“ (T. (1940) (meaercis]_1xtrbers

of tic Fnited Stotes. 1st Ed., XXI-fFfié 39., 312 fi3s.

:JZGTRJ-Vill thflc 30. 14M; , Jew \Tuflrf London.

Cblclrick, 1;. C. (-Axuxtccfl 'Ffitxyt of :311thotfix: :romtfi: sub-

stnizxtes; Fri if c yrm tmQ 01‘ cpitixink3s (if r.o<ml;7 ornio7norfi s].

glopts. (unpubliswed)

Lsrlinj, C. h. (1996) Sex 1L dioecious plorts. Full.

.ocpe” B 7. Club 33: 177-199.

---——~------—- (131?) Litosis in liviv3 03112. “W17

TOfirey Pot. Club 39: 407-409.

--———------——- (1923) Cbxomosome DGFQVIOP in Acev3§le+.r-

(“lilos L. g"fiéfl!‘.Cf”. JFUL". 39073. 10: 4.50-5.37.

first, K. L. (194L) 8 9 distribution OF "elf-Ltevilitg {r

the £10et'iJ'; {71-311111}. 17:70". 71917. “Lil. L-joc. 8?: (”/9-

519.

Foster, R. C. (1933) ChPOWOSOWS DHWFBP in A33? end Steph—

flec. Jour. 1:2: 1:“ . 14: Sit-393.

Freemqn, O. T. (1941) A red Tople, Silver maple Fybrid.

Jour. ERHWHilt; 32: ll—lf.

3 isor, L. G. (19?0) 9 list of clnomosome numbers in Au—

1 TQFWS. Geieiica U: (”l-464.

——--—----—--— (1939-0) A list of chromosome numbers in

3““i()s perwns. 13ib33<93r. ficovfi;. O: lVlmwfioo.

—------————-- (1930-9) A lisu Of cbfiowosome numbers in

KH,10~7c7ms. G ' 2 61—260.CD

:31

.DGD

“WCT‘

H C)

..D

H

7“ V" Ir 1:

1111131“. , 3.. o If”. Q "7! 1‘81‘P\ . - 1941) mcxtbook of dendrol-

7.777 57.136 Pd XV+ 5A9 foo €1.24.03- fies. McGraw-Hill Rock

00-, InC. Y W York & LO]

mes of some dioecious

Lindssv, R. H. (1950) The stroloso

2nii.osoerrxs Amer. Jknu7.'?ot. 17: ILm2-174.

Ksude, P. r. (1959) The h,rtou cata103ue. A list of tte

chromosome nU7c:919 of species of Priti31 flowering

plent . New 9*"tol. 39: l-S .

Laugmfln’ 0. Chronocowo moepholovy, constic doublibg “no

seconflnr" assCC1ctnon in Ate~ waiencifies L. Ho7editss

15: 145-175. 1933.

-----~--—~- (1933) nowc fiddltlhn”l rewewk" to tbc question

of‘ uolxrnlcixi acin“ filminnpvfix"os Triotsflfles. Lzrnor. 1%0c.

”WJLr ~10“. HHJA. _<: V2477? Citel f~om Plant Breed.

ulfl) Kitosis in the n.11en mother cells

1 " l

, o

of floor Pegundo J. eno “‘nobylco lVl in L. Ann. Hot.

("fi\ vs; ll,*- 1 :7 . A “““’”“‘“"'

:4”7s7“, '?. S. 5 Prwwvi ‘H. P. (1932) ’ 0" JWllfil till ers

0? Indie. filk-urttw, GOVerWHWn7t of Iz‘is Ceilfi7“l 3tjlic-

onion “hinca.

ijtrfiflxizii, E. 5;. ( $34) (Eilwlifitfi7t8 111 self-{<fllins—

. 13:10“. Oj‘a I ‘3, .2, iv -: {13(3‘373 t: ,13. ‘1p )0ijQ . jfi‘ (1qu :1 1:11.81). (no t . ;\(.‘{ (3 .

Sci. 9.9.9.”. 4: Q9”- )lo. (Cited f“0u Plsrt Domed.

Abstr. Vol 5, 19C5. )

-151-

Q ", \ ' x

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(\_ ~ 1 . (x ‘ ____ I ‘ ‘1 ’ 7‘ 71 1'!

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