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J. Japan. Soc. Hort. Sci. 54(1) : 46-57. 1985.
Overcoming the Self-incompatibility of Raphanus
sativus by Application of Amino Acids,
Vitamines and Phytohormonesl
Sachlko MATSUBARA
Faculty of Agriculture, Okayama University, T sushima, Okayama 700
Summary
Overcoming self-incompatibility of Raphanus sativus cultivars. `Motohashi-taibyo Minowase' (M-Mino) and `Minowase' (Mino) was studied by application of 15 different kinds of amino acids, two kinds of vitamines, four kinds of phytohormones and three kinds of phenolic substances. Effects differed between the two cultivars. In "M -Mino" , arginine (100 mg!l), serine and tyrosine (500 mg/l), alanine, valine and cystine (1000 mg/1), leucine (100 and 1000 mg/l), lysine, threonine and phenylalanine
(500 and 1000 mg/l) were significantly effective. On the other hand, in "Mino", arginine and leucine (200 mg/l) and histidine (1000 mg/l) were significantly effective. Phytohormones and vitamines tended to increase some fruit development and setting, although rate of them were not statistically significant. Besides, these chemicals in-duced some parthenocarpic fruits.
Introduction
Overcoming the self-incompatibility is one of the important step to obtain pure lines in incompatible species. This has been partially realized in some species of the family Cruci f erae by bud and delayed pollinations (1, 10, 31) and by several methods to treat stigma and pollen. Those treatments included cut-ting of stigma (33), washing of stigma sur-face or pollen with organic solvents (22, 23, 26, 37), mechanical and electric methods (27, 28), carbon dioxide treatment (3, 4,17,18), low and high temperature treatments (5, 6, 8, 12, 15, 29), treatment with relatively high hu-midity (21), mentor pollen method (7, 9, 24, 25, 34, 35) and treatments with phytohormone
(2, 16, 32), amino acid and vitamine (16). In the amino acid treatment, seeds were ob-tained with glutamic acid and glycine in self-incompatible mating combination of Rapha-nus sativus. Among vitamines, those were obtained with folic acid and nicotinic acid. Amino acids and phenolics are reported to be
')Received for publication September 18, 1981.
deeply related with sexual reproduction of
plants (11, 13, 14, 16, 30, 36). In this paper will report results of study
of effects of applications of 15 different kinds of amino acid selected from their promoting effect on plant growth, four kinds of phyto-hormone and three kinds of phenolic sub-stances for overcoming the self-incompatibili-ty of Raphanus sativus.
Materials and methods
Two cultivars of Japanese radish, Rapha-nus sativus cv. `Motohashi-taibyo Minowase'
(M-Mino) and cv. `Minowase' (Mino) were used as materials. Since plants of these cul-tivars were self-incompatible of sporophytic type, fruit setting is seldom obtained by pol-linating flowers within plant, while high fer-tility is seen by the cross-pollination between
plants which have different compatible geno-types. In the present study, for obtaining flowers as long period as possible, seeds were sown twice to soil inside a plastic house, in September, 1982 and in March, 1983, and
plants therefore flowered during a period
46
OVERCOMING THE SELF-INCOMPATIBILITY OF RAPNANUS SATIVUS 47
from April to June, 1983. Besides, some
young seedlings vernalized under low temper-ature at 5°C for 22 days were grown under
growth chamber conditions that temperature was controlled at 10°C with 16 hrs of day-length for 40 days. Plants with the latter treatment started to flower from September to November. Flowers which had been covered with paper bags one day before blooming for avoiding natural pollination by insects were treated with chemicals just after dissecting petals and filaments at their middle, and then self-pol-linated with pollen from anthers of the same flower. Pistils and part of petals and fila-ments left on flower were dipped into the solution used for the treatment for a few sec-onds. The solution contained singly follow-ing amino acids, vitamines, phytohormones or phenolic substances. Nine kinds of amino acid, glutamine, glycine, alanine, aspartic acid, serine, lysine, threonine, arginine and leucine and two kinds of vitamine, folic acid and nicotinic acid were singly dissolved at 50, 100 and 200 mg/l (hereafter indicated as low concentrations) and 500 and 1000 mg/l (high concentrations). Six amino acids, proline,
phenylalanine, histidine, cystine, valine and tyrosine were singly dissolved at the high concentrations. Four phytohormones, naph-thalene acetic acid (NAA), indole acetic acid
(IAA), kinetin and Ns-benzyl adenine (BA), were singly dissolved at 10 mg/l and low con-centrations. The phenolic substances, chlo-rogenic acid, caffeic acid and quercetin were singly dissolved at 10 mg/i and low concen-trations in 10% ethanol aqueous solution. Control pistils were dipped in water or 10%
ethanol aqueous solution before self- and cross-pollination. At about two months af-ter the pollination, number of fruits obtained from the treated flowers and number of seeds for each fruit was recorded. Difference of the effect by chemicals was calculated as ratio of fruit setting and seeds per fruit. For a statistical treatment Duncan's F-test was ap-
plied.
Results
All of 457 flowers made cross pollination set normal fruits in the experiment regardless the difference of cultivars and seasons, indi-cating that three different cultivation condi-tions did not disturb any in their reproduc-tion. On the other hand, by pollinating a flower with its self pollen, as expected, dras-tic reduction of number of fruit developed was resulted. As is shown in Table 1 for a total of 457 flowers pollinated in both culti-vars, 20 (4.4%) normal fruits were obtained. It does not to be any difference of this rate between two cultivars, for the rate were fair-ly constant despite number of flowers polli-nated were different. However, the rate seems to fluctuate to some extent depending on the cultivation seasons, although it is not very high. These indicate that the cultivars are genetically almost identical in their na-ture to produce fruit by self-pollination and therefore that results from two cultivars are able to pool together and variation by envir-onment conditions have to be considered in analyzing the effect of chemicals. In the case of the pollination after treating with chemicals, increased number of fruit setting as compared with those in the controls was
Table 1. Number of fruits obtained by self-pollination of Raphanus sativus cvs. M-Minowase and Minowase.
48 SACHIKO MATSUBARA
a
C)
0
U ro
0 C;
ro
0
C; 0
ro U
a a ro
C)
ro
ro
0 C)
0 C)
ro
C)
ro
0 C)
i
U
1--I
CC C) 'NN V
CC CC N V
V
C) .N
C) 0
ro C) .N
0 a
1"^q1 W
V̂, W.y C
.N ro
0
C)
w
N C) 0 ro H
OVERCOMING THE SELF-INCOMPATIBILITY OF RA PHANUS SATIVUS 49
v
O a
3 0
0
v
E C C a, a .
ii: 4. o
~ q o a
C U zaa
N >+ '*'
50 SACHIKO MATSUBARA
m a
x
C
U ro
C C
ro
0
C 0
ro U
a a
ro
v
ro
m ro
3 0
C
ro C
ro
ro
0 C
U
y C
C N 'Np V
C C C
C
C .N
C C ..w
ro C .N
0 a
V U
~..I C .H ro
O
V U
V/
N
ro H
OVERCOMING THE SELF-INCOMPATIBILITY OF RAPHANUS SATIVUS 51
0
0 a
N a a
0b
00
zz 0
N ?~
52 SACHIKO
resulted in some chemicals in both cultivars
(Table 2). The number of fruit per a flower pollinated clearly increased by threonine. So did in arginine except low concentrations of 50 and 100 mg/l and in leucine except 50 and 200 mg/l in plot of plants grown in 1982/9-1983/6 season. In the control of this season, the rate of fruit setting was further lower than those treated ; 8 fruits out of 313 polli-nated. Thus, increasing effect of the fruit setting by these three kinds of amino acids is thought to be indubitable. Among six kinds of amino acids tested only in one season of 1983/6-11, phenylalanine and cystine showed higher rate of fruit setting. With the rest of the 10 kinds result will be divided into four categories. One is these amino acids that the setting rate in low concentrations re-mained about the same level with the con-trol, and the second group of amino acid varied rate in concentration, probably also in season. And the amino acid in the third cate-
gory is one effect of which increased propor-tionally with the concentration of amino ac-ids. The fourth group of amino acid shows higher level of the fruit setting from lower concentrations however not increasing rate apparently in higher concentrations. In the
Table 4. Fruit obtained b
MATSUBARA
y self-pollination in Minowase after application of vitamine
first category include amino acids, glycine,
alanine and valine. For the second, gluta-
mine, aspartic acid, lysine, and probably pro-line and tyrosine. In the third only serine
is included. For the fourth, threonine, ar-
ginine, leucine, phenylalanine and cystein are included. For histidine the present data may not be big enough to determine the cat-
egory it belongs, although it might be of the third category.
In Table 3 is shown result for study on
number of seed per a fruit that consequently obtained from the fruits obtained by the
treatments. As is clear from the table, number of seeds obtained per fruit does not
show any difference that depends on the kind of treated amino acid. Total of 35 seeds
out of 20 fruits equalled 1.75 seeds per fruit in the control. Those in treated materials
seem to fluctuate around this value. How-
ever, it is interesting that a tendency is that higher the rate of fruit setting more the
number of seed per fruit. In the table is al-so shown the rate calculated for the total
number of flowers pollinated. As expected,
number of seeds accordingly decreased with total number of fruit obtained. In compari-
son with 0.08 seed obtained per total flowers
Raphanus sativus L, cvs. M-Minowase and on the pistil.
OVERCOMINGTHESELF-INCOMPATIBILITYOFRAPHANUSSATIVUS 53
incontrol,atotalof1,313seedsoutof2,965
flowerswithtreatmentequalsO.44seed,
showingthatthetreatmentasawholewas
effectiveinpartiallyovercomingtheself-in-
compatibility.Morethanoneseedperflow-
erwasobtainedinsometreatments.Espe一
ciallyhighernumberobtainedwas2.58seedsbytreatmentof500mg/lsolutionofarginine,
followedby2.36and2.09seedsbyserineand
asparticacid,respectively.Ontheotherhand,thetreatmentswithglutamineand
glycineremainedaboutthesamelevelwith
Table5.Seedobtainedbyself-pollinationinRaphanussativusL.cvs.
Minowaseafterapplicationofvitamineonthepistil .
M-Minowaseand
Cultivarandseason
ChemicalConcen-trationmg/1
M-Minowase1982/9-1983/6seed/seed/fruZflwY
1983/3-6seed/seed/
fruflw
Minowase1982/9-1983/6seed/seed/
fruflw
ユ983/3-6
seed/seed/
fruflw seed/fru
Total
seed/flw
folicacid
nicotinicacid
total
control
self-pollination
cross-pollination
50
100
200
500
1000
50
100
200
500
1000
2.2
4.5
2.0
0
4.0
2.0
1.6
4.6
0.23
0.59
0.50
0
0.08
0.04
0.04
4.60
2.5
3.1
0
∩)
3
21.6
4.6
0.33
1.65
0
0.23
0.16
4.60
0
1.0
1.5
2.0
2.0
2.3
3.0
3.6
0
0.06
0.09
0.12
0.11
0.27
0.08
3.60
1.0
1.5
0
0
3.5
3.6
o.07
0.21
∩)
0
0.21
3.60
11/5(2.2)
29/8(3.6)
25/13(1.9)
6/3(2.0)
31/11(2.8)
2/1(2.0)
6/2(3.0)
9/4(2.3)
0/0(0)
3/1(3.0)
122/48(2.5)
35/20(1.8)
11/70(0.16)
29/80(0.36)
25/77(0.32)
6/29(0.21)
31/31(1.00)
2/60(0.03)
6/73(0.08)
9/73(0.12)
0/75(0)
3/27(0.11)
122/595(0.21)
35/457(0.08)
1760/400(4.4)1760/400(4.40)
zNumberofseedperfruit .YNumberofseedperflowerpollinated .
Table6.Fruitobtainedbyself-pollinationinRaphanussativusL ,cvs.
Minowaseafterapplicationofphytohormoneonthepistil.
M-Minowaseand
Concen-trationmg/1
Cultivar
Chemical M-Minowase
fru/flwZ%Y
Minowase
fru/HwO fru/flw
Total
0
NAA
IAA
BA
Kinetin
total
control
self-pollination
cross-pollination
0
0
∩)
「0
0
0
1⊥
9白
0
0
0
FD
∩)
0
可⊥
∩∠
∩)
∩)
O
FO
∩)
0
1
9白
0
0
0
だ0
0
0
1
∩∠
2/34
4/40
4/42
12/39
16/40
16/32
8/31
7/41
2/35
7/31
8/34
1/35
7/68
70/70
5.9
ユ0.0
9.5
30.8
40.0
50.0
25.8
17.1
5.7
22.6
23.5
2.9
10.3
100.0
5/30
6/31
6/29
2/38
1/32
8/43
12/35
7/35
3/38
17/37
19/38
14/37
2/34
40/40
16.7
19.4
20.7
5.3
3.1
18.6
34.3
20.0
7.9
45.9
50.0
37.8
5.9
100.0
7/64
10/71
10/71
14/77
17/72
24/75
20/66
14/76
5/73
24/68
27/72
15/72
187/857
9/102
110/110
10.9
14.1
14.1
18.2
23.6
32.0
30.3
18.4
6.8
35.3
37.5
20.8
21.8
8.8
100.0
zNumberoffruitobtained/Numberoff' .owerpollinated.YPercentageoffruitobtained .
54 SACHIKO MATSUBARA
that in control. Result of the treatment with vitamines fo-lic acid and nicotinic acid is shown in Table 4. With folic acid fruit setting tended to be higher and the treatment with nicotinic acid did not increase any and tended to be even lower than the control. Similar tendency of increased number of seeds per a fruit is seen with folic acid, and with nicotinic acid the number remained to be in the same level throughout the different concentration
(Table 5). Table 6 shows result of the treatment with
some of phytohormones. Clear increase of number of fruits was obtained with kinetin. The rest of three kinds of phytohormones similarly tended to increase the setting rate. With BA the rate shows proportional reduc-tion with increasing concentration in both cultivars. A reverse tendency to BA is seen with IAA, and with NAA, except the rate is somewhat higher than in control any clear tendency with concentration is not able to find in the present result. As was in the rate of fruit setting, clear tendency to in-crease number of seeds per fruit was not found by the treatment with phytohormones
(Table 7). Besides amino acid and phytohormone, ef-
fect of phenolics was tested by similarly treat-ing pistils in their different concentration. Although with the treatment some fruits were obtained, the rate did not show any increase, not significantly different from the control statistically. Also those similarly treated with mixture solution of phytohor-mone and amino acid did not show any tendency of increasing rate. Accordingly, the data were not included in the tables.
Discussion
Some higher fruit setting in self-incom-
patible Raphanus cultivars was obtained in some of chemicals used in the present experi-ment. Among those three kinds of sub-stance phenolics did not show any apparent increase of the rate of number of fruit out of flowers pollinated by self-pollen after treatment of pistils. Apparent increase of the rate was obtained some of amino acids. Although that their physiological mechanism involved in overcoming self-incompatibility in which pollen tube germination and follow-ing elongation into pistil is biologically
Table 7. Seed obtained
Minowase after
by self-pollination in Raphanus sativus L.
application of phytohormone on the pistil.
cvs. M-Minowase and
OVERCOMING THE SELF-INCOMPATIBILITY OF RAPHANUS SAT V US 55
inhibited is at present not very clear, it has somehow indicated that phenolics do not affect any on that while some amino acids positively exerted to an extent to rise the fruit-setting rate distinctly higher than the range shown by the control pollination made in three different cultivation conditions. Generally the treatment with the concentration of 1000 mg/l showed higher rate of fruit setting. In lower concentrations, particular effect was obtained depending on the kind of amino ac-id. The effect of the fourth group of amino acid was particularly apparent from lower concentrations, but after reaching the certain level the effect tended to remain, rather sim-
ilar up to 200 mg/l, at 500 mg/l the effect was either increased twice or still remained at the same. This may indicate the presence of some other factor to inhibit or coun-teract toward the effect of the particular amino acid. On the other hand, effect of serine at 1000 mg/l is clearly nearing to the normal 100% of the cross-pollination. The nearly proportional increase of the rate from 0% at 50 mg/l to 72.7% at LOCO mg/l in-dicates the involvement of the concentration
depending reaction of really simple molecular level, in overcoming the self-incompatibility. Although this necessitates confirming the re-sult by repeating experiment, the mechanism to inhibit pollen tube germination, penetra-tion and its development by growing through style is overcome by the presence and involve-ment of serine molecules or some of com-
pound newly synthesized on pistil including serine. Certification of this assumption will have to be tested. Also there are possibilities that other amino acids that do not affect with
the concentrations in the present experiment may exert in the higher concentrations. At any rate that effect depends on concentration and that is seen in particular kinds of amino acids indicates its involvement in substantial
mechanism of self-incompatibility that might be a kind of the immunological reaction. The effectiveness of these amino acids, how- ever, is not clear to materials that do quietly not set fruit at self-pollination.
In overcoming the self-incompatibility, mechanical method to remove any disturb-
ance for pollen-tube growth or pistil has been reported by previous several authors. Those included simply removing stigma (33), washing surface of stigma with organic sol-vents (22, 23, 26, 37) and mechanical and elec-tric method that applied stigma (27, 28). All of these somehow directly removing or sof-ttening waxy substances that are present and covering the papillae. On the other hand, treatments with chemicals and with high temperature (5, 6, 8,12,15, 29) and CO2 (3, 4, 17,18) are thought to affect in altering the endogenous biochemical reaction in stigma. Amino acid may possibly involved as one of
such. It has reported that endogenous amino acid level of Nicotiana alata are different be-tween compatible and incompatible pollina-tion (11). The most marked changes were seen in an increase of r-aminobutyric acid and alanine and decrease of glutamic acid, and they were much greater following com-
patible pollination than incompatible one. Similar tendency of increase in compatible
pollination was also seen in the levels of valine, leucine and asparagine. Although these re-sults in tobacco, however, may be not direct-
ly related with the results of the present ex-
periment in radish, overcoming of self-in-compatibility may be induced with changes of endogenous level resulted by exogenous appli-cation of amino acids. Also reported is that S-allele specific proteins may differ by amino acid substitution (19, 20). Thus further study on the effect of amino acid would be interested and thought to be important not only overcoming self-incompatibility but also reaction when pollen germinate and grow on
pistils. Furthermore, although the applica-tion of amino acid may be a very practical method for overcoming self-incompatibility, it necessiates to compare with the effective-ness of bud pollination that has been used as a practical method.
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OVERCOMING THE SELF-INCOMPATIBILITY OF RAPHANUS SATVUS 57
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ア ミノ酸,ビ タ ミン及び植 物ホルモ ソ処理 に よる
ダイコンの自家不和合性 打破
松 原 幸 子
岡山大学農学部700岡 山市津島
摘
ダイ コンの2栽 培品種 「本橋系耐病美濃早生」 と 「美
濃早生」 の 自家不和合性打破 のため,15種 のア ミノ酸,
2種 の ビタ ミン,4種 の植物 ホルモン,3種 の フェ ノー
ル物質を用いて処理を行 った.処 理は各物質の水溶液に
雌ずいを浸漬後 自花受粉 を し,2か 月後結実率,種 子数
な どを調査 し,ダ ンカンのF検 定に よ り有意差の有無を
みた.
その結果,打 破の効果 は品種 と濃度に よ り差がみ られ
た.「 本橋系耐病美濃早生」ではア ルギ ニン(100mg/1),
セ リン,チ ロシン(500mg/1),ア ラニン,バ リン,シ
要
ス テ ィン(1000mg/1),ロ イ シ ン(100,1000mg/1),
リジ ン,ス レオ ニ ン,フ ェ ニル ア ラ ニ ン(500,1000mg/
1)が 有 効 で あ った.一 ・方 「美濃 早 生 」 は,ア ル ギ ニ ン,
ロイ シ ン(100mg/1),ヒ ス テ ィデ ィ ン(1000mg/1)が 有
効 で あ った.植 物 ホ ル モ ン,ビ タ ミンは 結 果 を 誘 導 し,
特 に 「本 橋 系 耐病 美 濃 早生 」 で はIAAが,「 美 濃 早 生 」
で は カイ ネ チ ンが 高 い 結 果 率 を 誘 導 した が,有 意差 は 認
め られ な か っ た.フ ェ ノー ル 類 は 効 果 が な か つ た.し か
しな が らいず れ の物 質 も,単 為 結 果 を 誘 導 した.
