GENERAL TECHNICAL REPORT

USDA FOREST SERVICEGENERAL TECHNICAL REPORT NC-53

Macronutrient deficiencysymptoms in seedlings of four

northern hardwoods

North Central Forest Experiment StationForest Service, U.S. Department of Agriculture

North Central Forest Experiment StationRobert A. Harm, Director

Forest Service - U.S. Department of Agriculture1992 Folwell Avenue

St. Paul, Minnesota 55108

Manuscript approved for publication August 23, 19781979

For sale by the Superintendent of Documents, U.S. Government Printing Office, Washington, B.C. 20402

Stock Number 001-001-00530-4

MACRONUTRIENT DEFICIENCY SYMPTOMS INSEEDLINGS OF FOUR NORTHERN HARDWOODS

Gayne G. Erdmann, Silviculturist,Frederick T. Metzger, Associate Silvicultu^

and Robert R. Oberg, Forestry ResearchMarquette, Michigan

Presently very little is known about the inor-ganic nutrient requirements of our importantnorthern hardwood species. Little informationconcerning optimum nutrient levels, criticalranges for essential elements, visual deficiencysymptoms, and physiological effects of nutrientdeficiencies has been published.

Detecting deficiencies is essential if timely cor-rective action is to be undertaken, A visual foliarsymptom approach is rapid, inexpensive, and reli-able as leaves are especially sensitive indicators ofdeficiencies (Kramer and Kozlowski 1960). How-ever, before any corrective action is taken, supple-mental chemical analyses of foliar tissue should bemade to verify suspected nutrient deficiencies(Phares and Finn 1972).

Deficiency symptoms for some hardwoods havebeen described: basswood (Ashby 1959, Ashby andMika 1959); pin oak (Hacskaylo and Struthers1959); locust and sweetgum (Hacskaylo 1960);American elm, box elder, hackberry, Russianolive, and silver maple (Sucoff 1961, Perala andSucoffl965, Lamb and Murphy 1968); eastern cot-tonwood (Hacskaylo and Vimmerstedt 1967); yel-low-poplar (Ike 1968); black locust, black walnut,eastern cottonwood and pin oak (Hacskaylo et al.1969); black walnut (Phares and Finn 1972); redmaple, gray birch, and black cherry (Walker1956); and red maple (Smith 1976). Except forAshby, Walker, and Smith, little information isavailable on the effects of deficiencies on thegrowth of northern hardwood species.

Several related studies have dealt with potas-sium deficiencies in American elm (Perala andSucoff 1965), effects of macronutrient and molyb-denum deficiencies on yellow birch growth and

nutrient uptake (LaFlamme and Lafond 1967,Hoyle 1971, and Hannah 1973), effects of variouslevels of N, P, and K on paper birch growth (Bjork-bom 1973), the relation of nitrogen deficiencies tosugar maple decline (Mader et al. 1969), and theeffect of Mn and Cu on P uptake in sugar maple(Gysietal. 1975).

Our studies were established to observe, photo-graph, and report distinctive visual deficiencysymptoms for the macronutrient elements (N, P,K, Ca, Mg, and S) in sugar maple (Acer saccharumMarsh.), red maple (Acer rubrum L.), white ash(Fraxinus americana L.), and paper birch (Betulapapyrifera Marsh.) seedlings. Deficiency symp-toms illustrated in this paper together withsupporting foliar analyses and growth data arepresented as preliminary guides in diagnosing nu-tritional disorders of these seedlings grown undernursery, greenhouse, and laboratory conditions.

The chemical composition and ovendry weightsof foliage, stems, and roots of seedlings grown incomplete single element deficiency solutions anddistilled water were compared. Seedling heightgrowth and the number of leaves produced perseedling under the various treatments were alsocompared.

METHODS

Sand culture nutrient deficiency studies withyoung sugar maple, red maple, paper birch, andwhite ash seedlings were conducted in the Forest-ry Sciences Laboratory greenhouse at Marquette,Michigan, The maples were studied in the summerof 1972; the ash and birch were studied in thesummer of 1973.

Sugar maple seeds with protruding radicles andemerging red maples in the cotyledon stage weregathered, rinsed thoroughly with distilled water,and planted directly in pots. Paper birch seedswere germinated in distilled water in petri dishesunder fluorescent light until cotyledons expanded.The radical end of white ash seed, which wasdifficult to germinate, was clipped to promote uni-form germination. After clipping, the seeds wereplaced on damp filter paper in petri dishes untilseeds with elongating radicles were available forplanting.

Two to three germinants were planted in 660grams of high-grade silica sand in 550 cm3 claypots lined with inert, clear, polyethylene bags. Pinholes were pricked into the plastic bags throughthe basal hole of the clay pot for bottom drainage.The sand in the 1973 birch and ash trials wasleached with 20 percent cold hydrochloric acid (for3 days followed by washing with distilled wateruntil there was no change in pH of the standingwater); unleached sand was used in the earlier1972 maple trials. The improved procedure in1973 was thought necessary to produce Fe defi-ciency symptoms1 and a better procedure forproducing Ca deficiency symptoms with the high-grade silica sand.

Germinants were grown in distilled water andthen conditioned in gradually strengthened com-plete nutrient solutions for 46, 25, 65, and 33 daysfor sugar maple, red maple, paper birch, and whiteash, respectively. Seedlings in each pot werethinned to leave the most vigorous individual.

Following the development of the first trueleaves, the seedlings were given a complete nutri-ent plus Fe-EDTA solution (Hacskaylo 1960) until120 ml of full strength solution per pot was accu-mulated and retained by capillary action. At theend of the conditioning period the pots wereflushed with 150 ml of distilled water and thedeficiency treatments were begun. Thereafter potswere reflushed with distilled H2O and replenishedwith appropriate fresh nutrient solutions at 2~week intervals.

^Results of the iron deficiency trials are not re-ported here because although visual symptomsdeveloped they could not be confirmed by the nutri-ent concentration percentages and there was nogrowth retardation in the paper birch or white ashseedlings.

Cultural solutions were complete except for theomission of one of six elements: N, P, K, Ca, Mg, orS. Treatments also included controls: one with thecomplete nutrient solution, and one with distilledwater only. Controls were used as checks on con-tamination and to provide a basis for judginggrowth responses after deficiency treatments wereapplied. Solutions were prepared with reagentgrade or high purity chemicals and pyrex-glassdistilled water (See Appendix, table 25 and 26 forcomposition, substitutions, and strength of the so-lutions used). Cultural solution pH values used inthe pots ranged from 4.8 to 5.5 and were 5.8 for thedistilled water. Watering and flushing kept pHvalues above pH 5.0 in all treatments. Moisturelosses were monitored with weighed representa-tive pots and replaced once or twice daily withdistilled water. Nutrient status in the pots wasmaintained by adding 10 ml of cultural solutiononce each week and completely flushing and re-plenishing with new nutrient solution every twoweeks. Injurious surface concentrations of salts,formed by rapid evaporation of cultural solutionsadded from above during hot weather (Hewitt1966), were avoided by top watering, breaking upsurface crusts weekly, and biweekly flushing.Maples were grown in deficient cultural solutionsfor 117 days; ash and birch for 113 days.

A randomized block design with 5 replications ofthe 6 deficiency and 2 control treatments was usedfor each species. Individual potted seedlingswithin each replication were the experimentalunits to which treatments were randomly applied.Each replication was formed by selecting seed-lings of uniform heights. There were two completenutrient controls in each birch and ash replicate,but only one for the maples.

Plants were grown under at least a 16-hourphotoperiod with supplemental fluorescent and in-candescent lighting of 200- to 300-foot candles atthe leaf surface. The greenhouse was covered witha shade cloth to reduce normal daylight intensityby 50 percent and to lower summer greenhousetemperatures. Day temperatures were kept below34C and night temperatures above 12C. Insectswere controlled by hand. Pots within replicationsand entire replications were rotated twice weeklyto equalize bench and lighting effects.

Visual deficiency symptoms were described asthey appeared, followed at weekly intervals

throughout their development, and photographedat significant stages. Symptoms included unusualcolor, necrosis, wilting, distortions; general sizeand growth of leaves, petioles, buds, and stems.Also noted were changes in color of interveinalareas, veins, bases, margins, and tips of young andold leaves. Types of necrosis in leaves included:marginal scorch, tip scorch, salting, spotting, andblotching. "Salting" refers to grain-sized spots ofsalt; "spotting" to pinhead-sized spots; and"blotching" to individual, usually irregularly-shaped areas larger than spots which became nec-rotic all at once. Significant growth reductions inseedling height growth and in ovendry weights ofcomponent parts as compared to the complete nu-trient controls were also considered diagnostic of asevere nutrient deficiency.

Seedling height growth was measured at 2-weekintervals. At completion of the study, leaves werecounted and plants were separated into leaves,stems, and roots. Chemical analyses of these com-ponents were made on composited samples fromthe five replications, ovendried at 70C, and groundin a Wiley2 mill to pass a 20-mesh screen. Variousstages of deficiency symptoms and leaf maturityare represented in the analyzed material. Totalnitrogen was determined by the semimicro-Kjel-dahl method (Bremner 1965); total S was by nitricperchloric acid digestion (Tabatabai and Bremner1970); and P, K, Ca, Mg, Fe, Al, Cu, Zn, B, and Mnby emission spectrography of dry-ashed (485C for4 to 5 hours) samples.3 Checks with duplicatesamples proved to be satisfactory.

Treatment effects on seedling growth, weight,and size were evaluated with an analysis of vari-ance and Duncan's New Multiple Range test. Com-positing samples to obtain sufficient material forchemical analyses ruled out statistical tests of theinfluence of treatments on nutrient content.

Supplemental laboratory studies were run inthe summer of 1975 to get better quality photos forP and K in sugar maple and P and S in white ashseedlings.

^Mention of trade names does not constituteendorsement of the products by the USDA ForestService,

^Chemical analysis were made by the ResearchAnalytical Laboratory, Department of Soil Science,University of Minnesota.

VISUAL DEFICIENCYSYMPTOMS, EFFECTS ONGROWTH AND CHEMICAL

COMPOSITION OFSEEDLINGS

Visual foliar symptoms of nutrient deficienciesare described separately for each species becausesymptoms for a given element often varied mark-edly among species. This approach should make ite*asier to pinpoint deficiencies within a species.Color photographs illustrate plants exhibiting dis-tinguishing foliar symptoms and advanced stagesof each deficiency.

Seedlings grown in complete nutrient solutionswere healthy plants with normal leaves (figs, 1A,2A, 3A, 4A). Their foliage was dark green exceptfor white ash which varied from green to darkgreen. Expanding apical leaves on all species nor-mally were light colored or reddish. These colorswere not indicative of deficiencies.

All seedlings grown in distilled water withoutnutrients were stunted and had small yellow-green, yellow, or red foliage and petioles (figs. IB,2B, 3B, 4B). Height growth on sugar maple and redmaple seedlings ceased immediately after theywere put in distilled water cultures, while whiteashes and paper birches stopped growing after 14and 21 days, respectively. Foliage on distilledwater seedlings was borne at abnormal angles tothe stem.

Besides foliar symptoms, many deficienciescaused obvious growth reductions and other ef-fects on the above-ground parts of seedlings. Thesegrowth losses often occurred before any visual fol-iar symptoms appeared, particularly in the casesof nitrogen, phosphorus, or calcium deficiencies.As with visual foliar symptoms, the impact of acertain nutrient deficiency on growth often dif-fered among species.

The effect deficiencies had on height growth var-ied with species. The height growth of white ashand sugar maple seedlings was not as greatlyaffected by deficiencies because they stoppedgrowing soon after treatments were begun. Paperbirch and red maple seedlings, on the other hand,continued growing longer, allowing more time forthe deficiencies to influence growth. (Only statisti-cally significant effects of seedling growth re-sponses are given in this section (tables 1-24).

Figure 1A.—Sugar maple complete nutrient control. Figure IB.—Sugar maple distilled water control.

Figure 2A.—Red maple complete nutrient control. Figure 2B.—Red maple distilled water control.

Figure 3B.—Paper birch distilled water control.

Figure 3 A.—Paper birch complete nutrient control.

Figure 4A.—White ash complete nutrient control.

Figure 4B.—White ash distilled water control.

Chemical analysis of seedling parts verified thedeficiencies, while the data provide a guide to themagnitude of the differences that can be expectedbetween abnormal and healthy seedlings else-

where. Nutrient content in stems, roots, and allfoliage of seedlings is expressed as the percentcomposition (concentration) and the total weight(total or absolute amount) of the element taken up.

SUGAR MAPLE

Nitrogen

Deficiency symptoms

Initially, entire leaf surfaces throughout theplant gradually changed from dark green to lightgreen. Some leaf lobe tips turned yellow and subse-quently scorched. After this the lower pair ofleaves turned completely yellow to contrast dis-tinctly with light-green upper leaves. In advancedstages this yellowing in lower leaves was followedby more necrotic tip and marginal scorching andextensive blotching which gradually progressedup the stem.

Growth

Shoot elongation almost stopped as soon as seed-lings were switched from complete to N-deficientcultural solution. As a result, they appearedstunted and had fewer and smaller leaves thancomplete nutrient control seedlings. Omitting Ncaused significant losses in total seedling dryweight (table 1) and all component parts (foliage,stem, roots) of it. N-deficient seedlings weighed 74percent less than the complete nutrient controlsand were comparable to the distilled water con-trols except root weights were greater for N-defici-ent seedlings.

Nutrient content

Total uptake in deficient seedlings was a fourthof that in complete controls. Concentrations invarious deficient seedling tissues fell to near 1percent (table 1).

Table 1.— Average height growth, ovendry seedling weight, nitrogen concentrations and total content incomponent parts of sugar maple seedlings grown in deficient, complete nutrient, and distilled watersolutions for 117 days1

Distinguishing symptoms (top), advancedsymptoms (bottom) in sugar maple.

Nitrogen

Solution

CompleteDeficientWater

Heightgrowth2

Mm27.43.02.2

Dryweight3Grams4.321.731.04

ConcentrationLeaves Stem

Percent ovendry2.240.820.43

1.881.20_4

Rootsweight

1.1.0.

LeavesTotal Content

StemMilligrams per

831717

323,1

702085

8.272.16

—

Rootsseedling44.2913.573.55

Plant

85.2618.93

—

'See Appendix table 28 for more complete growth data."Height growth from treatment establishment until harvest.30vendry weight of entire plant."Dashes indicate that inadequate material was available for chemical analysis.

Phosphorus

Deficiency symptoms

First, all leaves throughout the plant quicklydeveloped distinctive dull dark-green surfaces;light-green and yellow-green mottles and necrotictips showed up shortly thereafter. Later themottles became yellow and necrosis spread alongthe leaf margins. Finally, discolored, water-soaked-appearing spots developed on interveinaltissues of the lower leaves. These spots enlargedand rapidly coalesced into larger blotches. Spot-ting symptoms progressed upward from the lowerleaves.

Growth

Phosphorus, like N, was crucial to growth; itsomission caused stunting and fewer leaves wereproduced. All aspects of seedling growth were com-parable to growth of the distilled water controls.

Nutrient content

P-deficient seedlings took up 9 times less P andconcentrations were a third of normal (table 2).


Table 2.—Average height growth, ovendry seedling weight, phosphorus concentrations and total content incomponent parts of sugar maple seedlings grown in deficient, complete nutrient, and distilled watersolutions for 117 days1

Phosphorus

Solution


Heightgrowth2

Mm27.43.22.2

Dryweight3

Grams4.321.461.04

ConcentrationLeaves Stem Roots

Percent ovendry weight0.16 0.24 0.240.05 0.09 0.070.05 0.09 0.10

Total ContentLeaves Stem Roots Plant

Milligrams per seedling2.34 1.06 5.81 9.210.22 0.16 0.59 0.970.22 0.10 0.50 0.82

'See Appendix table 28 for more complete growth data.2Height growth from treatment establishment until harvest.30vendry weight of entire plant.

Potassium

Deficiency symptoms

Yellow leaf tips, petiole drooping, and necroticspotting were the earliest signs of K-deficiency.Symptoms developed slowly, beginning with lowerleaf margins turning yellow green and then gradu-ally yellow. Yellowing widened inward betweenthe green midrib and main veins, giving the re-maining green tissues the appearance of a deeplylobed white oak leaf. Similar symptoms appearedlater in upper leaves, but the green-patterned edgewas fringed with a purple tint. Finally, yellowareas bronzed or scorched and leaf margins curledup. Leaf petioles were also purple. Dead leavesremained attached to the stem.

Growth

Dry weight for all component parts were re-duced by omitting K, but stem and root weightswere reduced more than foliage weights. Yields inboth stems and roots in K-deficient seedlings werecomparable to those in the distilled water controls.

Nutrient content

Deficient seedlings showed 8 times less uptakeand lower concentrations of K, especially in theleaves and roots, than normal seedlings (table 3).


Table 3.—Average height growth, ovendry seedling weight, potassium concentrations and total content incomponent parts of sugar maple seedlings grown in deficient, complete nutrient, and distilled watersolutions for 117 days1

Potassium

Solution


Heightgrowth2

Mm27.410.62.2

Dryweight3

Grams4.321.651.04




Milligrams per seedling13.14 2.51 22.75 38.40

1.96 0.59 2.11 4.660.99 0.53 2.05 3.57

1See Appendix table 28 for more complete growth data.2Height growth from treatment establishment until harvest.30vendry weight of entire plant.

Calcium

Deficiency symptoms

Newly formed upper leaves were greatly re-duced in size by Ca deficiency. Overall fading fromlight- to yellow-green occurred first in upperleaves and then in lower leaves. Yellowing begansimultaneously in all upper leaf tips and prog-ressed inward toward the leaf base. Yellow leaftips turned bright red before dying while the restof the leaf remained yellow green. In more ad-vanced stages, the first or lowest pair of true leavesturned yellow before drying and rolling up.

Growth

Leaf number and all other aspects of seedlinggrowth were greatly diminished by the lack of Ca.Calcium-deficient seedlings were similar to thedistilled water control seedlings and weighed lessthan those grown under other deficiency treat-ments. Calcium-deficient seedlings were about 72percent lighter than the controls.

Nutrient content

Almost twelve times more Ca was taken up bynormal seedlings than those grown without Ca(table 4). Foliar and stem concentrations indicatedonly a threefold difference and root concentrationswere lower.


Table 4.—Average height growth, ovendry seedling weight, calcium concentrations and total content incomponent parts of sugar maple seedlings grown in deficient, complete nutrient, and distilled watersolutions for 117 days1

Calcium

Solution


Heightgrowth2

Mm27.44.62.2






3.96 0.67 0.61 5.224.17 0.92 1.10 6.19


Magnesium

Deficiency symptoms

The upper leaves developed a distinctive pat-tern: interveinal tissues turned light green whilemidribs, main veins, and adjacent tissues re-mained dark green. Leaf tip and edge yellowingand scorch also accompanied this symptom. Light-green mottles and yellow-green patches began ap-pearing on lower leaves next, but the distinctivemargin-to-rib pattern did not develop. Necrosis inold leaves consisted of scattered salting and spot-ting. In young leaves necrosis finally expandeddown between the main veins, and the dried leaftips curled down.

Growth

Dry weights of component parts of Mg-deficientseedlings declined, but seedling height growth,leaf numbers, and leaf size were not greatly af-fected by this deficiency.

Nutrient content

Total uptake by deficient seedlings was muchlower (10 times), especially in the roots (16 times)than complete control seedlings (table 5).


Table 5.—Average height growth, ovendry seedling weight, magnesium concentrations and total content incomponent parts of sugar maple seedlings grown in deficient, complete nutrient, and distilled watersolutions for 117 days*

Magnesium

Solution


Heightgrowth2

Mm27.417.42.2






'See Appendix table 28 for more complete growth data."Height growth from treatment establishment until harvest.30vendry weight of entire plant.

10

Sulfur

Deficiency symptoms

First signs of S deficiency appeared on new up-per foliage. These leaves were light green with apinkish cast, cupped downward and curled, andhad scorched tips; later they turned yellow greenand developed scorched margins. Lower leaves re-mained dark green at first, then gradually light-ened to a uniform yellow-green color with necrotictips and interveinal blotches.

Growth

Sulfur deficiency caused a moderate reductionin seedling height growth. Weights of deficientseedlings parts were significantly below those ofthe complete nutrient controls but yet almosttwice as heavy as the distilled water controls.

Nutrient content

Sulfur uptake was reduced about sixfold for thevarious seedling parts (table 6).


Table 6.—Average height growth, ovendry seedling weight, sulfur concentrations and total content in compo-nent parts of sugar maple seedlings grown in deficient, complete nutrient, and distilled watersolutions for 117 days1

Sulfur

Solution


Heightgrowth2

Mm27.48.62.2

Dryweight3Grams

4.322.261.04


Percent ovendry weight0.19 0.15 0.260.07 0.06 0.07_4


Milligrams per seedling2.77 0.66 6.29 9.720.49 0.14 0.92 1.55

— — — —

'See Appendix table 28 for more complete growth data.2Height growth from treatment establishment until harvest.30vendry weight of entire plant."Dashes indicate that inadequate material was available for chemical analysis.

11

RED MAPLE

Nitrogen

Deficiency symptomsNitrogen deficiency first showed up as a gradual

fading of the lower leaves from green to lightergreen in red maple seedlings. This fading symp-tom gradually progressed up the stem to the upperfoliage. In more advanced stages the lower foliageturned pink, then red, and the basal leavesscorched on the tip, turned brown, rolled up, andfinally abscised.

Growth

N-deficient seedlings weighed about half asmuch as the complete nutrient controls; most ofthe weight loss occurred in the foliage and roots.Newly formed leaves stayed noticeably smallerthan older leaves as the deficiency developed.

Nutrient content

Total amount of N in deficient seedlings wasabout one-fourth of normal but concentrationswere only about one-half of normal (table 7).

Distinguishing symptoms (top), advancedsymptoms (bottom) in red maple.

Table 7.—Average height growth, ovendry seedling weight, nitrogen concentrations and total content incomponent parts of red maple seedlings grown in deficient, complete nutrient, and distilled watersolutions for 117 days1

Nitrogen

Solution


Heightgrowth2

Mm154.4163.0

1.0


ConcentrationLeaves

10

StemPercent ovendry

.43 0.95

.71 0.374 —

Rootsweight

1.310.600.50

Leaves

256

Total ContentStem

Milligrams per.45 10.8374 2.85

—

Rootsseedling38.12

7.62—

Plant

74.4017.21

—


12

Phosphorus

Deficiency symptoms

Brown tip scorch and blotching in the lowerleaves were the first signals of P deficiency. Thenlower leaves turned dull green as upper leavesgradually became mottled dull green, yellow, andred. As the deficiency advanced young leaf tipsoften scorched while lower leaves developed dulllight-green, yellow-green, and red mottles, andmarginal scorch. Lower leaves browned com-pletely before they were finally shed.

Growth

Phosphorus-deficient seedlings were stunted,like the distilled water seedlings, and muchshorter than any other deficient seedlings. Phos-phorus-deficient seedlings had fewer, smaller, andnarrower leaves than normal seedlings. Totaloven dry weights of P-deficient seedlings wereidentical to the distilled water controls and aboutone-tenth that of normal seedlings.

Nutrient content

Absolute amount of P in deficient plants wasabout a fortieth of normal, yet concentrationsranged from about a half to a sixth of normal (table8).


Table 8.—Average height growth, ovendry seedling weight, phosphorus concentrations and total content incomponent parts of red maple seedlings grown in deficient, complete nutrient, and distilled watersolutions for 117 days1

Phosphorus

Solution


Heightgrowth2

Mm154.4

22.61.0







Potassium

Deficiency symptomsInitially, young foliage became lighter yellow

green, except for the veinal network which re-mained a contrasting darker green. Young leaftips also curled down and then scorched. Marginalscorching of young foliage followed, along withsalting in both young and old foliage. Salted areason young foliage coalesced into blotches as lowerfoliage turned light green and began to roll up.Finally lower leaves shriveled up, but they wereretained on the stem hanging down from horizon-tal petioles.

Growth

Potassium-deficient seedlings had shorter, moreslender stems with smaller leaves than normalseedlings. Dry weights of K-deficient seedlingswere comparable to distilled water seedlings.However, K deficiency reduced root and stemyields more than foliage yields.

Nutrient content

Total content of K taken up in tissues of defici-ent seedlings was reduced by a factor of 8 or morebut concentrations were only 2 to 5 times less(table 9).


Table 9.—Average height growth, ovendry seedling weight, potassium concentrations and total content incomponent parts of red maple seedlings grown in deficient, complete nutrient, and distilled watersolutions for 117 days1

Potassium

Solution


Heightgrowth2

Mm154.4

60.21.0

Dryweight3Grams

5.831.450.50





1.29 0.48 1.26 3.030.32 0.32 1.13 1.77


14

Calcium

Deficiency symptoms

First, young foliage faded from dark green tolight green. Then interveinal tissues turned yel-lowish green while midribs and veins stayed lightgreen. Next symptoms were the curling down ofyoung leaf tips and the appearance of red tips andmargins on older foliage. This was followed bynecrotic or discolored spotting on leaves of all ages.As the deficiency intensified, foliage from the bot-tom up the plant changed to first pink, then red,petioles drooped below horizontal, and leaf bladeshung vertically.

Growth

Calcium deficiency caused stunting and a greatreduction in leaf numbers and their size. Seedlingcomponent weight yields were not different thanthe distilled water controls.

Nutrient content

Deficient plants had 21 times less Ca in theirtissues than normal plants. The amounts of Ca inroot tissues were especially low. Although per-centage values showed similar trends, none of thevalues approached the reductions shown by abso-lute amounts (table 10).


Table 10.—Average height growth, ovendry seedling weight, calcium concentrations and total content incomponent parts of red maple seedlings grown in deficient, complete nutrient, and distilled watersolutions for 117 days1

Calcium

Solution


Heightgrowth2

Mm154.4

37.21.0






2.60 0.51 0.43 3.541.10 0.92 0.78 2.80


15


Magnesium

Deficiency symptomsInitially, Mg-deficient seedlings seemed more

luxuriant than seedlings receiving all nutrients.The first characteristic symptom appeared as aprominent pattern in upper leaves. This patternformed as interveinal tissues faded from the mar-gins inward between veins toward the midrib,leaving a broad, sharp zone of dark-green tissuesnext to the main veins and midrib. The secondcharacteristic symptom was the deep-purple colorthat developed in mature leaves as they becamedominated by anthocyanin pigments. This colora-tion progressed up the plant, while young leavesrolled up and developed tip scorch. After this, nec-rotic blotching intensified in older leaves just be-fore they were shed from the bottom up the stem.

GrowthMagnesium deficiency reduced seedling root

weights below the complete nutrient control levelbut foliage and stem weights were not. During therelatively short duration of these studies, the goodheight growth of red maple, white ash, and paperbirch seedlings in Mg-deficient treatments may becaused by small amounts of Mg remaining in thesand as contaminants, or perhaps by the effect ofsodium initially partly replacing .these species'needs for Mg. These effects under potassium defi-ciency are well known (Hewitt 1966).

Nutrient contentUptake of Mg was reduced considerably. Either

the percentage values or the total amount of Mg inthe foliage appear to be good diagnostic tools forestimating a seedling's Mg status (table 11).

Table 11.—Average height growth, ovendry seedling weight, magnesium concentrations and total content incomponent parts of red maple seedlings grown in deficient, complete nutrient, and distilled watersolutions for 117 days1

Magnesium

Solution


Heightgrowth2

Mm154.4222.0

1.0

Dryweight3Grams

5.834.550.50





0.85 0.20 0.70 1.750.31 0.08 0.30 0.69


16

Sulfur

Deficiency symptomsFirst, upper expanding leaf surfaces developed a

pink to yellow-green cast and their tips curledunder and drooped. Then tip scorch and spottingappeared on these leaves, which was later followedby the spreading of necrotic areas, first along themargins and then inward toward the midrib andpetiole. Old lower leaves turned lighter green thannormal and finally developed tip scorch.

Growth

Sulfur-deficient seedlings weighed about half asmuch as the complete nutrient controls after 117days under treatment. Besides this, the deficiencyalso noticeably depressed height growth and re-duced the size of younger leaves.

Nutrient content

Total amounts of S in deficient seedlings wasreduced sixfold, while concentrations indicatedabout half as much of a reduction from normallevels (table 12).


Table 12.—Average height growth, ovendry seedling weight, sulfur concentrations and total content incomponent parts of red maple seedlings grown in deficient, complete nutrient, and distilled watersolutions for 117 days1

Sulfur

Solution


Heightgrowth2

Mm154.4

79.21.0



Percent ovendry weight0.21 0.08 0.170.06 0.05 0.05

4



1See Appendix table 29 for more complete growth data.2Height growth from treatment establishment until harvest.

30vendry weight of entire plant."Dashes indicate that inadequate material was available for chemical analysis.

17

WHITE ASH

Distinguishing symptoms (top), advancedsymptoms (bottom) in white ash.

Nitrogen

Deficiency symptoms

Leaves and petioles throughout the white ashplants gradually faded to a uniform light-greencolor as N became deficient. Following this, thecotelydons and the lowest pair of true leavesquickly changed from light green to yellow greento entirely yellow, while all higher leaves retainedtheir light-green color. This is the characteristicsymptom for N deficiency in most plants. As greenleaves faded, leaf margins and interveinal tissuesoften developed a purplish cast but no distinctivepatterns were formed. Shortly thereafter, thesesymptoms began moving up the stem. Necrosisfirst appeared on the margins of the yellow cotely-dons and subsequently as tip and marginal scorchon the lower leaves. Petioles also became moreerect than normal, having about a 15° angle withthe stem.

Growth

Nitrogen-deficient seedlings generally wereshorter and had smaller leaves than normal seed-lings. Deficient seedlings had the lowest total andcomponent weights of the 7 single-element defi-ciencies studied. Their weights were comparableto distilled water controls and 62 percent belowcomplete nutrient controls.

Nutrient content

Total uptake was about one-sixth of normal andconcentrations about half of normal in N-deficientseedlings (table 13).

Table 13.— Average height growth, ovendry seedling weight, nitrogen concentrations and total content incomponent parts of white ash seedlings grown in deficient, complete nutrient, and distilled watersolutions for 113 days1

Nitrogen

Solution


Heightgrowth2

Mm622839

Dryweight3Grams

3.511.331.11

ConcentrationLeaves

100

StemPercent ovendry32 1.10.54 0.77,40 0.66

Rootsweight

1.0.0.

185651

LeavesTotal Content

StemMilligrams per

9.11 5.611.73 1.540.96 1.12

Rootsseedling27.144.543.57

Plant

41.867.815.65


Phosphorus


Deficiency symptoms

Although foliage might fade slightly at first andsome terminal leaves might scorch at the tip, ma-jor P-deficiency symptoms began in the lowerleaves and slowly progressed up the stem. Shortlyafter, lower leaves turned lighter green, and pur-plish spots appeared on them. Next they developedyellow and brown necrotic areas, while upperleaves remained green. Symptoms in the veinalnetwork of lower leaves developed more slowly;the network remained light green as the leaf yel-lowed and finally turned yellow as the leaf browns.

Growth

Normally, growth reductions would be expectedby omitting P, but white ash is a determinategrower and 60 percent of its height growth wasalready completed before deficiency treatmentswere begun. Another growing period would be re-quired for the expected reductions to develop.

Nutrient content

Total uptake in tissues of deficient seedlingswas one quarter of normal. The reduction in up-take was much greater in foliage than other tis-sues (table 14).

Table 14.—Average height growth, ovendry seedling weight, phosphorus concentrations and total contentin component parts of white ash seedlings grown in deficient, complete nutrient, and distilledwater solutions for 113 days1

Phosphorus

Solution


Heightgrowth2

Mm626839

Dryweight3Grams

3.513.141.11






19

Potassium

Deficiency symptoms

Upper leaves might droop from normally erectpetioles before the tips scorched on terminalleaves. This was followed by the light-green fadingof upper leaf margins. Finally, all leaf tissue be-came the same color, but leaf ribs and bases stayedgreen longer. These symptoms gradually moveddown the stem to the lowest leaves. In severestages of K deficiency, upper leaf tips and marginsoften became completely scorched resulting incurved-up and rolled-in upper leaf margins. Later,some tips and edges scorched in lower leaves too.

Growth

Height growth was not reduced as expected forthe reasons discussed under growth of P-deficientwhite ash seedlings.

Nutrient content

Absolute amounts of K in leaf and stem tissuesand K-leaf concentrations reflect the reduceduptake of K which averaged about one-third ofnormal (table 15).


Table 15. — Average height growth, ovendry seedling weight, potassium concentrations and total content incomponent parts of white ash seedlings grown in deficient, complete nutrient, and distilled watersolutions for 113 days1

Potassium

Solution


Heightgrowth2

Mm627939

Dryweight3Grams

3.513.591.11






Calcium

Deficiency symptoms

Early symptoms of Ca deficiency were tipscorching of terminal leaves, death of uncurlingnew leaves and growing tips, and erect petiolesthat almost parallel the stem. In extreme casespetioles of terminal leaf pairs crossed each other.Fading at first resulted in uniform light-greenleaves and petioles, but then intensified in inter-veinal areas to a very pale green to whitish greento a whitish yellow, with some areas almost white.Through the early interveinal color changestissues around leaf midribs and major veinsremained light green. In severe stages, water-soaked-looking blotches appeared on the edges,and the interveinal tissues scorched. Similarsymptoms appeared later on lower leaves.

Growth

Stems and leaves appeared smaller than normalon Ca-deficient plants. Root weights of deficientplants were especially low, being 60 percent belownormal levels and comparable to distilled watercontrols.

Nutrient content

Total Ca uptake was a sixth and concentrationsabout a fourth of normal levels (table 16).


Table 16.— Average height growth, ovendry seedling weight, calcium concentrations and total content incomponent parts of white ash seedlings grown in deficient, complete nutrient, and distilled watersolutions for 113 days1

Calcium

Solution


Heightgrowth2

Mm625639

Dryweight3Grams

3.511.791.11






21

Magnesium

Deficiency symptoms

Distinctive symptoms of Mg deficiency first ap-peared in the lower two pairs of leaves, but soonwere present even in the youngest foliage. Leaftips, margins, and interveinal areas first faded to alight green, then to a yellow-green, and finallyyellow dominated. Tissues next to the veinal net-work, midribs, and point of petiole attachmentremained green, contrasting with other yellowleaf tissues. A purplish discoloration often pre-ceded necrosis that began at leaf tips and laterappeared at leaf margins and between veins.

Growth

Root weights of Mg-deficient seedlings wereslightly lower (14 percent) than those of completenutrient controls. Usually, the aboveground por-tions of Mg-deficient seedlings looked healthy untilthe deficiency was well developed. (See discussionof growth of Mg-deficient red maple seedlings.)

Nutrient content

Total Mg uptake and concentrations were aboutone-fourth that of normal seedlings (table 17).


Table 17.—Average height growth, ovendry seedling weight, magnesium concentrations and total content incomponent parts of white ash seedlings grown in deficient, complete nutrient, and distilled watersolutions for 113 days}

Magnesium

Solution


Heightgrowth2

Mm62

11739







22

Sulfur

Deficiency symptoms

Characteristic symptoms of S deficiency inwhite ash first appeared when young upper leavesturned lighter green than lower leaves. Newlyemerging upper leaves often rolled under andwrinkled. Then, leaves throughout the plant be-came a uniform yellow-green color. Tip scorch andspotting also occurred simultaneously on the up-per and middle leaves of some plants. In moreadvanced stages new growing points often died,and upper leaves turned entirely yellow except fortips and other scorched spots.

Growth

Root weight was reduced (36 percent) by theomission of S. Other expected growth reductionsdid not occur as already explained under thegrowth section for P-deficient seedlings.

Nutrient content

Total uptake of S in deficient seedling was abouta third of normal (table 18).


Table 18.— Average height growth, ovendry seedling weight, sulfur concentrations and total content incomponent parts of white ash seedlings grown in deficient, complete nutrient, and distilled watersolutions for 113 days1

Solution

Sulfur

Height Drygrowth2 weight3

Concentration Total ContentLeaves Stem Roots Leaves Stem Roots Plant


Mm628339

Grams3.512.621.11

Percent ovendry weight0.150.06

4

0.070.05

-

0.090.04

-

Milligrams per seedling1.040.38

-

0.360.25

-

2.070.59

-

3.471.22

-


Distinguishing symptoms (top), advancedsymptoms (bottom) in paper birch.

PAPER BIRCH

NitrogenDeficiency symptoms

Foliage throughout N-deficient paper birchseedlings quickly faded to a uniform light-greencolor while midribs and veins on the lowest leavesturned red. Shortly thereafter, oldest (lower)leaves turned yellow but their midribs and veinsremained red. These symptoms gradually pro-gressed up the stem while the lower leaves oftentip scorched before they turned completely brownand abscised.

Growth

Omission of N caused the greatest growth sup-pression of the six elements studied. Stunted N-deficient plants looked like those plants grown indistilled water. Leaf production was drasticallycurtailed in N-deficient plants and leaves weresmaller in size. Dry weight yields of leaves, stems,and roots were reduced to only about 10 percent ofthe complete nutrient controls. (See Appendix ta-ble 31 for more information.)

Nutrient content

Uptake was drastically curtailed in N-deficientplants. Absolute amounts of N averaged one-six-teenth of normal, but concentrations ranged fromnearly equal in stems and roots to about half nor-mal levels in the foliage (table 19).

Table 19.— Average height growth, ovendry seedling weight, nitrogen concentrations and total content incomponent parts of paper birch seedlings grown in deficient, complete nutrient, and distilled watersolutions for 113 days'

Nitrogen

Solution


Heightgrowth2

Mm179

1710

Dryweight3Grams

4.590.400.35


Percent ovendry weight1.12 0.62 0.950.52 0.60 0.850.44 - 4 0.66


Milligrams per seedling18.48 4.09 21.66 44.230.88 0.30 1.53 2.710.62 1.06


Phosphorus

Deficiency symptoms

Visual foliar symptoms of P deficiency were slowin showing up. At first, all leaves turned darkergreen and duller than normal. After this, dulllight-green and later dull yellow-green areas ap-peared at the tips, serrated edges, and in irregularpatches on the lower leaves. Then, chlorotic leaftips, edges, and some blotches turned brown as thesymptoms spread across the leaf and began tomove up the stem. In later stages, brown leaf tipscurled up before the leaves became entirely brownand abscised.

Growth

Height growth was also restricted by the omis-sion of P, but not to the same extent as in N-deficient seedlings. Upper leaves on P-deficientseedlings were noticeably narrower in width thannormal leaves but not in length. Average dryweight yields of birch foliage, stems, and rootswere reduced 62, 47, and 60 percent by P deficien-cy, respectively.

Nutrient content

Total P uptake— especially in the foliage— wasgreatly reduced by its omission from the culturalsolution. Concentrations showed similar patternsbut their magnitude was not as great (table 20).


Table 20.—Average height growth, ovendry seedling weight, phosphorus concentrations and total content incomponent parts of paper birch seedlings grown in deficient, complete nutrient, and distilled watersolutions for 113 days1

Phosphorus

Solution


Heightgrowth2

Mm17910010

Dryweight3

Grams4.591.880.35






25

Potassium

Deficiency symptoms

First symptoms of K deficiency appeared in theupper leaves before any color change in the lowerleaves took place. The upper leaves on some plantsbuckled under along the midribs and curved underat the margins. Then lower leaf tips, serrations,margins, and finally interveinal tissues, from themargins toward the midribs, gradually began toturn from light green to yellow green to yellow.This was followed by leaf tip and edge scorch in thelower leaves.

Growth

After growing in K-deficient solutions for 113days, birch seedlings exhibited only slight differ-ences in growth from normal seedlings.

Nutrient content

Total content of K was about a third of normal indeficient plants. Percentage values in tissuesshowed similar decreases (table 21).


Table 21.— Average height growth, ovendry seedling weight, potassium concentrations and total content incomponent parts of paper birch seedlings grown in deficient, complete nutrient, and distilled watersolutions for 113 days1

Solution

Potassium

Height Drygrowth2 weight3

Concentration Total ContentLeaves Stem Roots Leaves Stem Roots Plant


Mm179171

10

Grams4.594.780.35

Percent ovendry weight1.460.391.14

0.400.160.36

0.520.200.33

Milligrams per seedling24.09

6.281.60

2.641.200.18

11.864.860.53

38.5912.34

2.31


26

Calcium

Deficiency symptomsAll foliage slowly faded to a uniform light green

with basal leaves often appearing even paler. In-terveinal areas on the lower one or two leavesfinally turned pale yellow; narrow bands of lightgreen remained next to the veins and midribs.After yellowing, the tips and serrated edges oflower leaves scorched; water-soaked blotchessometimes appeared on them. Another advancedsymptom was the "hooking down" of young leaftips.

Occasionally veins in basal leaves appearedmuch like those on N-deficient seedlings. How-ever, in Ca-deficient seedlings the colored veinsymptom occurred much later, was not as sharplydefined, and appeared more as a reddish or pinkishcast around the veinal network.

Growth

Calcium deficient seedlings had fewer leavesand were about half as tall as normal seedlings.Deficient seedling weights averaged 47 percentbelow controls; 52 percent reductions occurred inboth stems and roots and a 39 percent reduction inthe foliage.

Nutrient content

Absolute uptake of Ca in tissues was a tenth to atwentieth of normal, but concentrations were onlya fifth to a tenth of normal (table 22).


Table 22.— Average height growth, ovendry seedling weight, calcium concentrations and total content incomponent parts of paper birch seedlings grown in deficient, complete nutrient, and distilled watersolutions for 113 days1

Calcium

Solution


Heightgrowth2

Mm1796910

Dryweight3Grams

4.592.430.35





3.33 0.35 1.54 5.221.67 0.58 0.85 3.10


27

Magnesium

Deficiency symptoms

A very distinctive chlorotic pattern developedon all but the youngest leaves in Mg-deficient pa-per birch. Tips, margins, and tissues betweenveins on mature lower and middle leaves firstbecame light green, while broad bands of tissueadjacent to midribs and main veins, at the base,and sometimes at the tip remained dark green.Light-green margins and interveinal tissues grad-ually changed to yellow green, then almost white.Some mature leaf tips scorched early, then, aftertissues had lightened considerably, necrotic spot-ting and blotching advanced rapidly.

Growth

Omission of Mg inhibited birch root and foliagegrowth more than stem growth. Root weights, af-ter 113 days under treatment, were only half ofnormal, while total weights were about 34 percentbelow normal levels. (See discussion of growth ofMg-deficient red maple seedlings.)

Nutrient content

The greatest reduction of Mg uptak e from nor-mal occurred in the leaves; the entire plant took upone-twelfth of normal amounts. Deficient tissueconcentrations ran from one-fourth to one-four-teenth of normal (table 23).


Table 23.—Average height growth, ovendry seedling weight, magnesium concentrations and total content incomponent parts of paper birch seedlings grown in deficient, complete nutrient, and distilled watersolutions for 113 days1

Magnesium

Solution


Heightgrowth2

Mm179191

10

Dryweight3

Grams4.593.020.35






Sulfur

Deficiency symptoms

Curling down and scorching of newly emergingupper leaf tips and edge serrations often were thefirst visible signs of S deficiency. New paper birchleaves, developing after deficiencies began, weremuch more yellow-green than the older foliagewhich remained dark green, except for yellowingof leaf tips and serrations. Later, old foliage fadedto the same color as the new foliage, but leaf tipsand margins were either yellow or scorched. Final-ly, necrotic spotting and edge blotching appearedin the oldest foliage.

Growth

Sulfur-deficient seedlings were slightly smallerthan normal and the development of new foliageappeared especially limited. Sulfur deficiency,like Mg deficiency, restricted root and foliage de-velopment more than stem development. Totalseedling dry weights were reduced 37 percent be-low normal levels by this deficiency.

Nutrient content

S uptake was reduced primarily in the foliage(table 24).


Table 24.— Average height growth, ovendry seedling weight, sulfur concentrations and total content incomponent parts of paper birch seedlings grown in deficient, complete nutrient, and distilled watersolutions for 113 days1

Sulfur

Solution


Heightgrowth2

Mm179128

10



Percent ovendry weight0.22 0.06 0.040.05 0.04 0.05

4



'See Appendix table 31 for more complete growth data.2Height growth from treatment establishment until harvest.30vendry weight of entire plant.'Dashes indicate that inadequate material was available for chemical analysis.

29

DISCUSSION

Visual foliar symptoms of nutrient deficienciesshould not be used alone for diagnosis becausedeficiencies of one element may be difficult todistinguish from another element and they canresemble damage caused by insects, disease, cli-mate, and pollution. Smith (1976) reported thatleaf symptoms of green veins with yellow interve-inal tissue in red maple, are similar for manganeseand iron deficiency and are often confused. Accord-ing to Worswick (1950), micronutrient deficienciesof manganese and copper are difficult to distin-guish from K-deficiency in fruit trees. Marginalleaf and tip browning caused by high temperature,leaf scorch following hot drying winds, and fluo-ride injury may all resemble K deficiency (U.S.Department of Agriculture, Forest Service 1973).

Visual deficiency symptoms usually will not beas well defined in nature as in our experimentsbecause they may be the result of interactions ormay be confounded by other deficiency or toxicitysymptoms. Thus, visual foliar symptoms can bethe first clue to which element might be limitinggrowth, but chemical analysis should be used toconfirm the suspected deficiency.

Samples from the foliage are normally analyzed,but sometimes other parts provide a better criticaldiagnosis (Hewitt 1966). Often a chemical soilanalysis of available nutrients is made, in additionto the other diagnostic measurements, to give amore complete picture of the nutrient status of thetrees in nurseries and in field fertilization pro-grams. Foliar analysis for trees is reviewed byLeaf (1973) and van den Driessche (1974). Theseprocedures are not yet routine and the results ofchemical analysis of soils or plant tissue must beinterpreted and applied carefully.

In our study, all leaves, regardless of maturity,were harvested to make chemical analyses. Theseedlings we analyzed generally showed moderateto severe deficiency symptoms. Any seedlings atthese deficiency levels would be under severe nu-trient stress so foliar symptoms would be veryapparent. When symptoms are not so clear orwhen growth is being restricted making a differen-tial diagnosis requires careful sampling, chemicalanalysis of the foliage, and expert advice (Pharesand Finn 1972). Common sense is required in mak-ing a correct diagnosis too, to assure that frost,drought, and diseases (Phares and Finn 1972) or

air pollution (U.S. Department of Agriculture,Forest Service 1973) are not causing the suspectednutrient problem.

Users wishing to make comparisons with ourdata for determining whether a nutrient deficien-cy exists, should collect all the leaves from a ran-dom sample of 10 seedlings in a suspected bed orunit after 115 days or in mid-August. Make surethe leaves are clean and avoid collecting themwithin several days after rain because of nutrientleaching (Phares and Finn 1972). The samples,including leaves and petioles, should be collectedin paper bags and air dried.

Compare the laboratory test results with our"complete" and "deficient" levels. Any levels at orabove our "complete" nutrient control levels (ex-cept possibly N which should be somewhat higherand is discussed below) are probably within arange of concentrations satisfactory for growth.Levels approaching our "deficient" levels wouldlimit growth.

Results of these chemical analyses are applicableto seedlings grown under greenhouse, laboratory,and nursery conditions, but not to forest-grownseedlings or those under field conditions where onlymature leaves are normally collected and analyzedand where leaf maturity, leaf position, and season-al variation in mineral absorption are taken intoaccount in sampling the nutrient content of foliage.

Critical deficiency, toxicity, and adequate nutri-ent supply levels have not been established formost of the northern hardwood species. However,Bjorkbom (1973) reported tentative optimum lev-els of 3 to 4 percent of foliar N as being adequate forbirch seedlings. His results indicate that our Nlevel was below that required for optimum seed-ling growth. Other leads from field studies, al-though not directly comparable, suggest that thefoliar levels of N in our complete nutrient controlswere not optimum for seedling growth. From fieldstudies, Mitchell and Chandler (1939) estimated"optimum" foliar N levels for mature sugar mapletrees to range from 2.77 to 2.85 percent. Our com-plete nutrient control foliar N level of 2.24 percentis within their adequate "working region" of 1.75to 2.77 percent N. Their region where N was limit-ing growth was below 1.75 percent, Mader et al.(1969) considered foliar N levels below 1.50 per-cent important in sugar maple decline. Mitchelland Chandler (1939) also estimated optimum and

30

other critical leaf N concentrations for red mapleand white ash trees. Our foliar N levels of 1.43percent N for red maple and 1.32 percent N forwhite ash seedlings (complete nutrient solution)are beginning to limit growth.

In western Massachusetts, Mader et al. (1969)associated sugar maple decline with N deficiency,described foliar symptoms typical of N deficiency,and showed a positive response to N fertilization inaffected trees.

Bjorkbom (1973) found that paper birch seed-lings were not greatly affected by different levelsof phosphorus or potassium. In New York, Walker(1956) described foliar symptoms of K deficiencyoccurring in red maple growing on abandonedfarms in K-deficient glacial outwash sands.Walker (1956) considered about 0.60 percent foliarK in mid-August as the point where deficiencysymptoms would be expected to appear under fieldconditions.

Besides the clues from field studies, tables ofnatural foliar nutrient concentrations have beenpublished that may also be used as guides in for-ests (Gerloff, Moore, and Curtis 1964, Henry1973).

Practical methods for correcting nutrient defi-ciencies are available from agriculture and vari-ous forest studies (Beaton 1973) but very few fromfertilization trials of northern hardwoods. In gen-eral, success in correcting deficiencies has beenachieved with both soil applications and foliarsprays of the appropriate fertilizers. This is a rap-idly expanding field and the reader is encouragedto consult the various testing agencies (Au-chmoody and Filip 1973) for advice. Foliar and soilchemical analyses can be purchased from stateextension services and universities.

LITERATURE CITEDAshby, William C. 1959. Limitation to growth of

basswood from mineral nutrient deficiencies.Bot. Gaz. 121(l):22-28.

Ashby, William C., and Edward S. Mika. 1959.Sulfur deficiency in Tilia americana. Bot. Gaz.

Auchmoody, L. R., and S. M. Filip. 1973. Forestfertilization in the eastern United States:hardwoods. In Forest fertilization Symp. Proc. p.211-225. U.S. Dep. Agric. For. Serv., Gen. Tech.

Rep. NE-3, 246 p. U.S. Dep. Agric. For. Serv.,Northeast. For. Exp. Stn., Broomall, PA.

Beaton, J. D. 1973. Fertilizer methods and applica-tions to forestry practice. In Forest fertilizationSymp. Proc. p. 55-71. U.S. Dep. Agric. For. Serv.,Gen. Tech. Rep. NE-3. U.S. Dep. Agric. For.Serv., Northeast. For. Exp. Stn., Broomall, PA.

Bjorkbom, John C. 1973. Response of paper birchseedlings to nitrogen, phosphorus, and potas-sium. U.S. Dep. Agric. For. Serv., Res. Note NE-157, 4 p. U.S. Dep. Agric. For. Serv., Northeast,For. Exp. Stn., Broomall, PA.

Bremner, J. M. 1965. Total nitrogen.In Methods ofSoil Analysis, Part 2. p. 1149-1178. Am. Soc.Agron., Inc., Madison, WI.

Gerloff, G. C., D. G. Moore, and J. T. Curtis. 1964.Mineral content of native plants of Wisconsin.Univ. Wisconsin Exp. Stn. Res. Rep. 14, 27 p.Madison, WI.

Gysi, C., C. H. Winget, and B. Bernier. 1975. Inter-actions of P, Mn and Cu in nutrient uptake bysugar maple. Can. J. For. Res. 5(1): 105-108.

Hacskaylo, John. 1960. Deficiency symptoms inforest trees. Trans., 7th Int. Congr. Soil Sci. Vol.Ill: 393-405, Madison, WI.

Hacskaylo, John, R. F. Finn, and J. P. Vimmer-stedt. 1969. Deficiency symptoms of some foresttrees. Ohio Agric. Res. and Dev. Cent. Res. Bull.1015, 68 p. Wooster, OH.

Hacskaylo, John, and P. Struthers. 1959. Correc-tion of lime-induced chlorosis in pin oak. OhioAgric. Exp. Stn. Res. Circ. 71,5 p., Wooster, OH.

Hacskaylo, John, and J. P. Vimmerstedt. 1967.Appearance and chemical composition of eas-tern cottonwood grown under nutrient deficientconditions. Ohio Agric. Res. and Dev. Cent. Res.Bull. 1004, 19 p. Wooster, OH.

Hannah, Peter R. 1973. Phosphorus stimulatesgrowth of yellow birch seedlings. Tree PlantersNotes 24(1):1, 2, 11.

Henry, Douglas G. 1973. Foliar nutrient concen-trations of some Minnesota forest species. Univ.Minnesota For. Res. Note 241,4 p. St. Paul, MN.

Hewitt, E. J. 1966. Sand and water cultural meth-ods used in the study of plant nutrition. Com-monwealth Agricultural Bureaux. (Revised 2ndEdition).

Hoyle, M. C. 1971. Effects of the chemical environ-ment on yellow-birch root development and topgrowth. Plant and Soil 35(3):623-633.

31

Ike, Albert F. 1968. Symptoms of nutrient defi-ciency in yellow-poplar seedlings. U.S. Dep.Agric. For. Serv., Res. Note SE-94,4 p. U.S. Dep.Agric. For. Serv., Southeast. For. Exp. Stn.,Asheville, NC.

Kramer, Paul J., and Theodore T. Kozlowski.1960. Physiology of trees. McGraw-Hill Co.,Inc., New York. 642 p.

LaFlamme, Yvon, and Andre Lafond. 1967. (Theinfluence of molybdenum on the development ofsome forest tree species). Contrib. Fonds. Rech.For. Univ. Laval 12, 19 p. Quebec, Canada.

Lamb, Fred M., and Wayne K. Murphy. 1968. Thegrowth and anatomical features of nutrient-de-ficient seedlings. In Eighth Lake States ForestTree Improv. Conf. Proc., p. 48-51. U.S. Dep.Agric. For. Serv., Res. Pap. NC-23, 60 p. U.S.Dep. Agric, For. Serv., North Cent. For. Exp.Stn., St. Paul, MN.

Leaf, Albert L. 1973. Plant analysis as an aid infertilizing forests. In Soil testing and plant anal-ysis. Walsh, Leo M. and James D. Beaton, eds.p. 427-454. Soil Sci. Soc. Am., Inc., Madison, WI.

Mader, Donald L., Bruce W. Thompson, and Jan P.Wells. 1969. Influence of nitrogen on sugar ma-ple decline. Massachusetts Agric. Exp. Stn.Bull. 582, 19 p. Univ. Massachusetts, Amherst.

Mitchell, Harold L., and Robert F. Chandler, Jr.1939. The nitrogen nutrition and growth of cer-tain deciduous trees of northeastern UnitedStates. Bull. 11, 94 p. Black Rock Forest, Corn-wall-on-the-Hudson, NY.

Perala, Donald A., and Edward Sucoff. 1965. Diag-nosing potassium deficiency in American elm,silver maple, Russian olive, hackberry and boxelder. For. Sci. ll(3):347-352.

Phares, Robert E., and Raymond F. Finn. 1972.Using foliage analysis to help diagnose nutrientdeficiencies in black walnut. North. Nut Grow.Assoc. Ann. Rep. 62:98-104.

Smith, Elton M. 1976. Manganese deficiency-com-mon in maples. Am. Nurseryman 143(1):11,131, 132.

Sucoff, Edward I. 1961. Diagnosing nitrogen defi-ciency in silver maple. Univ. Minnesota For.Note 108, 2 p. St. Paul, MN.

Tabatai, M. A., and J. M. Bremner. 1970. A simpleturbidimetric method of determining total sul-fur in plant materials. Agron. J. 62(6):805-806.

U.S. Department of Agriculture, Forest Service.1973. Air pollution damages trees. 32 p. North-east. Area. State and Private Forestry, UpperDarby, PA.

van den Driessche, R. 1974. Prediction of mineralnutrient status of trees by foliar analysis. TheBot. Rev. 40(3):347-394.

Walker, Lawrence C. 1956. Foliage symptoms asindicators of potassium-deficient soils. For. Sci,2(2):113-120.

Worswick, G. D. 1950. Tree symptoms and leafanalysis determine potash needs. Better Cropswith Plant Food 34(ll):19-22, 41-43.

32

APPENDIX

Table 25.— Composition of the complete nutrientsolution

MACRONUTRIENTS

Compound or salt MolarityKN03

Ca(N03)2-4H20MgS04-7H20KH2P04

0.0020.0030.0020.002

MICRONUTRIENTS

FE-EDTAH3B03

MnCI2-4H20ZnCI2CuCI2-2H20Mo03

0.00008910.00003740.00000700.00000070.00000030.0000003

Table 26.—Composition of the substitute salt solutions for deficient cultures showing solutions to omit andto replace

Deficiency Omit Replace with-N-P-K~Ca-Mg-S

KN03, Ca(N03)2'4H20KH2P04

KN03, KH2P04

Ca(N03)2-4H20, MgS04'7H20MgS04'7H20MgS04-7H20

0.002M KCI, 0.003M CaCI2'6H200.002M KCI

0.002M NaH2P04, 0.002M NaN03

0.002M Na2S04, 0.003M Mg(N03)2-6H200.002M Na2S04

0.002M MgCI2-6H20

33

Table 27.—Average nutrient concentrations in foliage, stems, and roots of complete nutrient control seedlings

FOLIAGESpecies N P K

Percent ovendrySugar mapleRed mapleWhite ashPaper birch

2.241.431.321.12

0.16.17.36.51

0.90.78

1.421.46

Ca

weight2.382.241.311.87

Mg

0.43.63.25

1.17

S

0.19.21.15.22

Fe

129127194252

Zn

14.512.516.29.6

Cu

p. p.m.2.62.23.02.0

Mn

19.314.843.248.0

B

43.744.924.765.3

Al

10075

10474

STEMS

Sugar mapleRed mapleWhite ashPaper birch

1.88.95

1.10.62

.24

.15

.14

.19

.57

.36

.78

.40

1.431.12

.641.10

.16

.12

.14

.12

.15

.08

.07

.06

32403052

11.58.05.5

12.8

2.82.7

.73.2

29.86.7

19.310.4

21.910.119.212.2

27201128

ROOTS

Sugar mapleRed mapleWhite ashPaper birch

1.831.311.18

.95

.24

.25

.19

.21

.94

.59

.88

.52

.76,73.40.68

.30

.19

.13

.14

.26

.17

.09

.04

21120050

448

33.022.414,6

209.4

7.210.24.0

99.9

8.56,47.1

12.0

8.85.3

13.49.4

18831749

1578

Table 28.—Average height growth, stem height, number of leaves, ovendry weight of foliage, stem,and root of sugar maple seedlings after 117 days under deficiency treatments

Treatment

CompleteD-H20-N-P-K-Ca-Mg-S

Heightgrowth1

mm21 A ab2

2 .2c3.0 c3.2 c

10.6 abc4.6 c

17.4 abc8.6 be

Height3

100 a78 a74 a74 a80 a81 a91 a84 a

Number ofleaves

12.0 a7. 2 be6.0 c6.6 be9.0 abc6.8 be8.2 abc

10.0 ab

Average dry weight per seedlingFoliage

1.46 a0.43 c0.39 c0,44 c0.85 b0.46 c0.89b0.70 be

Stem

0.44 a0.11 d0.18cd0.18 cd0.18 cd0.16 cd0.19cd0.24 be

Root

gms2.42 a0.50e1.16 bed0.84 cde0.62 de0.61 de0.66de1.32 be

Total

4.32 a1.04d1.73cd1 .46 cd1 .65 cd1.23d1 .74 cd2.26 be

'Height growth is the difference in heights between the time deficiency treatments were begun and harvest."Values followed by the same letter in a column are not significantly different (0.05 level) using Duncan's New Multiple Range Test.3Total seedling height measured from the root collar to the top of the terminal bud.

34

Table 29,—Average height growth, stem height, number of leaves, ovendry weight of foliage, stem, and root ofred maple seedlings after 117 days under deficiency treatments

Treatment

CompleteD-H.,0-N-P-K-Ca-Mg-S

Heightgrowth1

mm154.4 ab2

1.0d163.0 a

22.6 cd60.2 cd37.2 cd

222.0 a79. 2 c

Height3

227 ab77 d

232 a91 cd

136 cd115 cd297 a153 cd

Number ofleaves

19.2 ab7.6 d.

18. Sab9.6 cd

14.4 be12.8 cd22.0 a12.6cd


1.78 a0.13d0.95 b0.17cd0.76 bed0.45 bed2. 13 a0.80 be

Stem

1.14aO. IOde0.77 abc0.09e0.24 de0.27 cde1.01 ab0.60 bed

Root

.gms2.91 a0.27c1.27 be0.26 c0.45 c0.71 be1.41 b1.62 b

Total

5. 83 a0.50d2.99 be0.52 d1 .45 cd1 .47 cd4.55 ab3.02 be

'Height growth is the difference in heights between the time deficiency treatments were begun and harvest.2Values followed by the same letter in a column are not significantly different (0.05 level) using Duncan's New Multiple Range Test,3Total seedling height measured from the root collar to the top of the terminal bud.

Table 30.—Average height growth, stem height, number of leaves, ovendry weight of foliage, stem, and root ofwhite ash seedlings after 113 days under deficiency treatments

Treatment


Heightgrowth1

mm62 bed2

39 cd28 d68 be79 b56 bed

117a83 ab

Height3

156 bed120 cd128cd167 abed168 abc149 bed209 a194 ab

Number ofleaves

10.4 ab7.2c8.4 be9.6 be

10.6 ab9.4 be

12.8 a10.8 ab


0.69 a0.24 b0.32 b0.57 a0.70 a0.49 ab0.82 a0.64 a

Stem

0.51 a0.17 b0.20 b0.50 a0.55 a0.39 ab0.62 a0.50 a

Root

gms2.30 a0.70 c0.81 c2.07 a2.35 a0.91 be1.97ab1 .48 abc

Total

3.51 ab1.11 cd1.33d3.14ab3.59 a1.79 bed3.41 ab2. 62 abed

'Height growth is the difference in heights between the time deficiency treatments were begun and harvest.2Values followed by the same letter in a column are not significantly different (0.05 level) using Duncan's New Multiple Range Test.Total seedling height measured from the root collar to the top of the terminal bud.

35

Table 31.— Average height growth, stem height, number of leaves, ovendry weight of foliage, stem, and rootof paper birch seedlings after 113 days under deficiency treatments

Treatment


Height

growth1

179ab2

10 e17e

100 cd171 ab69 d

191 a128 be

Height3mm

202 ab35 e42 e

125cd193 ab95 d

213 a155 be

Number of

leaves

25.5 ab9.4 d8.8 d

21 .6 be31. 6 c17.6 c34.2 a26.6 ab

Average dry weight per seedling

Foliage

1.65b.14f.17f.62 e

1.61 be1.01 de1 .38 cd1.15d

Stem

.66 b

.05 d

.05 d

.35 c

.75b

.32 c

.47 be

.55 be

Root

.gms2.28 b

.16 d

.18d

.91 c2.43 b1.10c1.17c1.18c

Total

4.59 b.35d.40 d

1.88c4.78 b2.43 c3.02 c2.88 c

'Height growth is the difference in heights between the time deficiency treatments were begun and harvest.2Values followed by the same letter in a column are not significantly different (0.05 level) using Duncan's New Multiple Range Test.3Total seedling height measured from the root collar to the top of the terminal bud.

36

Erdmann, Gayne G., Frederick T. Metzger, and Robert R. Oberg.1979. Macronutrient deficiency symptoms in seedlings of four northern

hardwoods. U.S. Dep. Agric. For. Serv,, Gen. Tech. Rep.NC-53, 36 p. U.S. Dep. Agric. For. Serv., North Cent. For. Exp. Stn.,St. Paul, MN.

Illustrates and describes the visual deficiency symptoms for N, P, K,Ca, Mg, and S in sugar maple, red maple, white ash, and paper birchseedlings. Effects of these deficiencies on the development and nutri-ent composition of seedlings are also examined.

OXFORD: 424.7. KEY WORDS: (Acer saccharum Marsh.), (Acerrubrum L.), (fraxinus americana L.), (Betula papyrifera Marsh.),chemical composition of tissues, dry-matter production, seedlingdevelopment.

Erdmann, Gayne G., Frederick T. Metzger, and Robert R. Oberg.1979. Macronutrient deficiency symptoms in seedlings of four northern

hardwoods, U.S. Dep. Agric. For. Serv., Gen. Tech. Rep.NC-53, 36 p. U.S. Dep. Agric. For. Serv., North Cent. For. Exp. Stn.,St. Paul, MN.

Illustrates and describes the visual deficiency symptoms for N, P, K,Ca, Mg, and S in sugar maple, red maple, white ash, and paper birchseedlings. Effects of these deficiencies on the development and nutri-ent composition of seedlings are also examined.

OXFORD: 424.7. KEY WORDS: (Acer saccharum Marsh.), (Acerrubrum L.), (Fraxinus americana L.), (Betula papyrifera Marsh.),chemical composition of tissues, dry-matter production, seedlingdevelopment.

Plant a tree...trees give oxygen.