新疆5个驼舌草二型花柱居群交互式雌雄异位的变异

doi:10.11983/CBB22225

摘要/Abstract

摘要： 异型花柱是被子植物的一种花柱多态现象, 表现为具有交互式雌雄异位的花型间传粉的亲和性, 以及花型内和自交的不亲和性。而同型花柱(homostyly)是在异型花柱形成与演化中出现的一种雌雄蕊同长且自交亲和的花型。驼舌草属为白花丹科中较早出现二型花柱的属, 野外调查发现分布于新疆的驼舌草(Goniolimon speciosum)自然居群出现大量雌雄蕊同长(H)的花型, 类似于同型花柱。该花型的产生背景是否已转变为型内和自交亲和, 它与典型的长(L)/短(S)花柱型花之间的亲和性关系等问题尚不清楚。该研究对驼舌草5个自然居群的花型构成与频率、花部特征参数和柱头花粉负荷进行统计, 并检测附属多态性和异型不亲和系统。结果表明, 所有居群均由L、S和H三种花型构成, 但在花粉纹饰和柱头表皮细胞形态上仍具二型性和严格的非选型交配, 表现为花粉纹饰和柱头乳突细胞形态不同的花型间亲和, 反之则不亲和。研究表明, H型花可能是由L型和S型花的雌雄蕊间距缩短所致, 是花型间交互式雌雄异位变异的结果, 但该花型并未转变为同型花柱。对于该物种而言, 这种交互式雌雄异位的变异与生理不亲和性的转变可能相对独立。

关键词: 驼舌草, 异型不亲和系统, 异型花柱, 同型花柱, 交互式雌雄异位

Abstract: Heterostyly is a kind of floral morph polymorphism in angiosperms, which is characterized by the reciprocal herkogamy and accompanied by physiological self- and intra-incompatibility but inter-compatibility. And the homostyly (H) is a kind of self-compatible floral morph with stigmas and anthers at the same position within a flower, which occurs during the formation and evolution of heterostyly. It has been reported that distyly appeared earlier in Goniolimon than in other genus in the Plumbaginaceae. However, our field survey found that a large number of floral morphs with the same length of pistils and stamens (H-morphs) appeared in the natural population of Goniolimon speciosum in Xinjiang, similar to homostyly. It is not clear how the H-morphs emerge, whether the H-morphs have been self- and intra-compatible, and what kind of relationship between the H-morphs and the typical floral morphs with long- (L-) or short- (S-) style flower. Thus, we investigated the floral morph composition and frequency, measured the floral parameters and stigmatic pollen loads among floral morphs, and also examined the accessory polymorphism and heteromorphic incompatibility system of G. speciosum in five natural populations. The result showed that all populations were composed of L-, S- and H-morphs, but each population still had dimorphism of pollen-stigma morphology and strictly disassortative mating, which showed that the floral morphs with different pollen ornamentation and stigma papilla cell morphology were compatible regardless of the reciprocal herkogamy, but incompatible if the floral morphs were the same. Conclusion: H-morphs may the result of the variation of reciprocal herkogamy between floral morphs by shortening of stigma in L-morph or shortening of anther in S-morphs, but have not transformed into the homostyly. For this species, the variation of reciprocal herkogamy and the transformation of physiological incompatibility may be independent of each other.

Key words: Goniolimon speciosum, heteromorpic incompatibility system, heterostyly, homostyly, reciprocal herkogamy

任登芙, 翟雅芯, 张爱勤. 新疆5个驼舌草二型花柱居群交互式雌雄异位的变异. 植物学报, 2023, 58(5): 733-742.

Ren Dengfu, Zhai Yaxin, Zhang Aiqin. The Variation of Reciprocal Herkogamy in Five Distylous Populations of Goniolimon speciosum in Xinjiang. Chinese Bulletin of Botany, 2023, 58(5): 733-742.

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https://www.chinbullbotany.com/CN/Y2023/V58/I5/733

图/表 5

图1 驼舌草的花枝、花型、花粉及柱头形态 (A) 花枝; (B), (C) S型和L型花; (D) H型花雌雄蕊; (E), (H) L型花的花粉粒和柱头形态; (F), (I) S型和HS型花的花粉粒和柱头形态; (G), (J) HL型花的花粉粒和柱头形态。S型花: 柱头显著低于花药的花, 即短花柱型花; L型花: 柱头显著高于花药的花, 即长花柱型花; H型花: 柱头和花药高度在同一位置的花, 即同长花柱型花; HS型花: 花粉纹饰和柱头表皮细胞形态与S型花一致的花; HL型花: 花粉纹饰和柱头表皮细胞形态与L型花一致的花。(A) Bar=2 cm; (B)-(D) Bars=2 mm; (E)-(G) Bars=10 μm; (H)-(J) Bars=20 μm

Figure 1 The flowering branch, floral morphs, pollen and stigma morphology in Goniolimon speciosum (A) Flowering branch; (B), (C) Flowers with S- and L-morph; (D) Pistil and stamen of H-morph; (E), (H) Pollen and stigma morphology of L-morph; (F), (I) Pollen and stigma morphology of S- and HS-morph; (G), (J) Pollen and stigma morphology of HL-morph. S-morph: Flower with stigma significantly lower than the anther, as short-styled morph flower; L-morph: Flower with stigma significantly higher than the anther, as long-styled morph flower; H-morph: Flower with stigma and anther in the same position, as homostylous morph flower; HS-morph: Flower with the same pollen and stigma morphology as S-morph flower; HL-morph: Flower with the same pollen and stigma morphology as L-morph flower. (A) Bar=2 cm; (B)-(D) Bars=2 mm; (E)-(G) Bars=10 μm; (H)-(J) Bars=20 μm

表1 5个驼舌草居群的花型构成与频率

Table 1 The floral morph composition and frequency in five populations of Goniolimon speciosum

Population	Location	Number of plant samples (L, S, H)	Floral morph frequency (L/S/H)	Proportion of H-morph (%)	χ²(L/S)	P-values (L/S)
KNS	87.024°E, 48.698°N, 1424 m	166, 212, 91	1.82/2.33/1	32.62	5.598	0.018
HXG	87.984°E, 43.811°N, 1853 m	104, 162, 105	0.99/1.54/1	28.30	12.674	0.000
NS	87.233°E, 43.517°N, 1904 m	82, 113, 44	1.86/2.57/1	18.41	4.928	0.026
GG	87.167°E, 43.500°N, 1919 m	44, 51, 34	1.29/1.50/1	26.36	0.516	0.473
ZS	80.988°E, 43.150°N, 1980 m	52, 68, 40	1.30/1.70/1	25.00	2.133	0.144

表2 驼舌草喀纳斯(KNS)居群的花部特征参数及3个自然居群各花型间柱头花粉负荷比较

Table 2 Comparisons of floral parameters of Kanasi (KNS) population and stigmatic pollen loads among floral morphs in three field populations in Goniolimon speciosum

Floral parameter	Population	S-morph (n=20-30)	L-morph (n=20-30)	H-morph (n=20-30)
Corolla tube length (mm)	KNS	(6.20±0.10)^a	(6.03±0.11)^a	(6.17±0.11)^a
Corolla opening diameter (mm)	KNS	(6.17±0.09)^a	(6.02±0.14)^a	(6.09±0.13)^a
Pistil height (mm)	KNS	(6.93±0.09)^c	(8.39±0.09)^a	(7.84±0.11)^b
Stamen height (mm)	KNS	(8.26±0.06)^a	(7.13±0.06)^c	(7.81±0.07)^b
Number of pollen grains per flower	KNS	(1127±46)^a	(1158±36)^a	(1110±24)^a
Number of ovules per flower	KNS	1	1	1
Number of stigmatic pollen loads (total, inter-morph)	KNS	(17±3)^b, (2.5±0.7)^a	(27±4)^ab, (1.4±0.4)^a	(33±7)^a, (1.3±0.5)^a
	GG	(25±3)^b, (0.6±0.3)^a	(49±3)^a, (1.1±0.3)^a	(56±13)^a, (1.6±0.9)^a
	ZS	(22±3)^b, (0.7±0.3)^a	(54±4)^a, (1.1±0.3)^a	(54±5)^a, (1.1±0.3)^a

图2 驼舌草喀纳斯(KNS)居群不同花型性器官高度的分布

Figure 2 The distribution of sexual organ height among different floral morphs of Kanasi (KNS) population in Goniolimon speciosum

图3 驼舌草4种授粉处理下结实率比较不同小写字母表示各处理间差异显著(P<0.05)。

Figure 3 Comparisons of fruit set of floral morphs under four pollination treatments in Goniolimon speciosum Different lowercase letters indicate significant differences among treatments (P<0.05).

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