Selection for reproductive traits and their adaptation in Stellera chamaejasme in degraded alpine grassland

doi:10.11686/cyxb2020182

Abstract

Abstract:

The reproductive traits of angiosperms are widely believed to have evolved largely through pollinator-mediated natural selection. The present study focused on Stellera chamaejasme, a dominant poisonous species in degraded alpine meadow in Tianzhu County of Gansu Province in China. S. chamaejasme was chosen as a model species to explore adaptation of floral traits and possible selective mechanisms by detecting their selection differentials and selection gradients. It was found that flower number per inflorescence and corolla tube length had both significant selection differentials and selection gradients (P<0.05), indicating positive directional selection for both traits. Also, significant nonlinear selection gradients (P<0.05) were detected for these two traits, possibly indicating stable selection occurring for them. Negative directional selection was detected for inflorescence length, with a significant linear selection gradient of -0.102±0.039 (P=0.008). No significant selection was detected for the other traits measured, such as corolla entrance size and inflorescence number per plant. In conclusion, the data indicate that some reproductive traits in S. chamaejasme exhibited a trend of adaptive evolution under natural selection, and individuals, which have relatively shorter inflorescence length, longer corolla tubes and more flowers per inflorescence, tend to be selected for due to their advantage in achieving reproductive fitness.

Key words: S. chamaejasme, reproductive traits, selection gradient, fitness, pollination

Wang-long LUO, Jian-qiang XIA, Jia-xin LI, Shu-fan SUN, Rui WANG, Bo ZHANG. Selection for reproductive traits and their adaptation in Stellera chamaejasme in degraded alpine grassland[J]. Acta Prataculturae Sinica, 2021, 30(4): 121-129.

Figures/Tables 5

References 46

1	Harder L D， Johnson S D. Darwin’s beautiful contrivances： Evolutionary and functional evidence for floral adaptation. New Phytologist， 2009， 183（3）： 530-545.
2	Campbell D R， Waser N M， Price M V. Mechanisms of hummingbird-mediated selection for flower width in Ipomopsis aggregata. Ecology， 1996， 77（5）： 1463-1472.
3	Stebbins G L. Adaptive Radiation of reproductive characteristics in angiosperms， I： Pollination mechanisms. Annual Review of Ecology and Systematics， 1970， 1（1）： 307-326.
4	Sletvold N， Grindeland J M， Gren J. Pollinator-mediated selection on floral display， spur length and flowering phenology in the deceptive orchid Dactylorhiza lapponica. New Phytologist， 2010， 188（2）： 385-392.
5	Benitez-Vieyra S， Glinos E， Medina A M N， et al. Temporal variation in the selection on floral traits in Cyclopogon elatus （Orchidaceae）. Evolutionary Ecology， 2012， 26（6）： 1451-1468.
6	Zhao Z G， Huang S Q. Differentiation of floral traits associated with pollinator preference in a generalist-pollinated Herb， Trollius ranunculoides （Ranunculaceae）. International Journal of Plant Sciences， 2013， 174（4）： 637-646.
7	Totland O. Environment-dependent pollen limitation and selection on floral traits in an alpine species. Ecology， 2001， 82（8）： 2233-2244.
8	Conner J K， Hartl D L. A primer of ecological genetics. Sinauer Associates Incorporated， 2004： 199-208.
9	Zhang B， Li Q J. Phenotypic selection on the staminal lever mechanism in Salvia digitaloides （Labiaceae）. Evolutionary Ecology， 2013， 28（2）： 373-386.
10	Gomez J M， Bosch J， Perfectti F， et al. Spatial variation in selection on corolla shape in a generalist plant is promoted by the preference patterns of its local pollinators. Proceedings of the Royal Society B： Biological Sciences， 2008， 275（1648）： 2241-2249.
11	Maad J， Alexandersson R. Variable selection in Platanthera bifolia （Orchidaceae）： Phenotypic selection differed between sex functions in a drought year. Journal of Evolutionary Biology， 2004， 17（3）： 642-650.
12	Sandring S， Rllhmaki M A， Savolainen O. Selection on flowering time and floral display in an alpine and a lowland population of Arabidopsis lyrata. Journal of Evolutionary Biology， 2007， 20（2）： 558-567.
13	Obeso J R. The costs of reproduction in plants. New Phytologist， 2002， 155（3）： 321-348.
14	Lu N N， Liu Z H， Ma Y， et al. Phenotypic selection analysis of flower traits in Delphinium kamaonense var. glabrescens （Ranunculaceae）. Biodiversity Science， 2019， 27（7）： 772-777.
	路宁娜，刘振恒，马妍，等. 展毛翠雀的花性状表型选择. 生物多样性， 2019， 27（7）： 772-777.
15	Zhao C Z， Gao F Y， Sheng Y P， et al. Fine-scale spatial distribution and spatial association of Stellera chamaejasme population. Arid Land Geography， 2011， 34（3）： 492-498.
	赵成章，高福元，盛亚萍，等. 狼毒种群小尺度空间分布格局及空间关联性研究. 干旱区地理， 2011， 34（3）： 492-498.
16	Gao F Y， Zhao C Z， Zhuo M， et al. Spatial distribution and spatial association of Stellera chamaejasme population in the different altitude in in degraded alpine grassland. Acta Ecologica Sinica， 2014， 34（3）： 605-612.
	高福元，赵成章，卓马，等. 高寒退化草地不同海拔梯度狼毒种群分布格局及空间关联性. 生态学报， 2014， 34（3）： 605-612.
17	Xing F， Wang Y H， Guo J X. Spatial distribution patterns and dispersal mechanisms of the seed population of Stellera chamaejasme on degraded grasslands in Inner Mongolia， China. Acta Ecologica Sinica， 2004， 24（1）： 143-148.
	邢福，王艳红，郭继勋. 内蒙古退化草原狼毒种子的种群分布格局与散布机制. 生态学报， 2004， 24（1）： 143-148.
18	Sun G， Luo P， Wu N， et al. Stellera chamaejasme L. increases soil N availability， turnover rates and microbial biomass in an alpine meadow ecosystem on the eastern Tibetan Plateau of China. Soil Biology and Biochemistry， 2009， 41（1）： 86-91.
19	Zhao C Z， Zhang Q P. The spatial pattern of soil seed bank of Stellera chamaejasme community in degraded grassland of the Qilian Mountains. Chinese Journal of Grassland， 2010， 32（1）： 79-85.
	赵成章，张起鹏. 祁连山退化草地狼毒群落土壤种子库的空间格局. 中国草地学报， 2010， 32（1）： 79-85.
20	Zhang Q， Zhao C Z， Dong X G， et al. Relationships between flower size， flower number， and plant size of Stellera chamaejasme population along an altitude gradient of degraded alpine grassland in Northwest China. Chinese Journal of Ecology， 2013， 32（12）： 3160-3166.
	张茜，赵成章，董小刚，等. 高寒退化草地不同海拔狼毒种群花大小，数量与个体大小的关系. 生态学杂志， 2013， 32（12）： 3160-3166.
21	Zhang Q， Zhao C Z， Dong X G， et al. Relationship between flower size and leaf size， number of Stellera chamaejasme population of degraded alpine grassland along an altitude gradient. Chinese Journal of Ecology， 2015， 34（1）： 40-46.
	张茜，赵成章，董小刚，等. 高寒退化草地不同海拔狼毒种群花大小与叶大小，叶数量的关系. 生态学杂志， 2015， 34（1）： 40-46.
22	Zhang Z Q， Zhang Y H， Sun H. The reproductive biology of Stellera chamaejasme （Thymelaeaceae）： A self-incompatible weed with specialized flowers. Flora-Morphology， Distribution， Functional Ecology of Plants， 2011， 206（6）： 567-574.
23	Zhang G， Guo Y Z， Zhang S P， et al. Harm status and control strategy of poisonous weeds of natural grasslands in Tianzhu County of Gansu Province. Progress in Veterinary Medicine， 2019， 40（3）： 123-128.
	张庚，郭亚洲，张水平，等. 甘肃天祝县天然草地毒草危害状况调查与防控. 动物医学进展， 2019， 40（3）： 123-128.
24	Shi Y， Hu T H， Gao H J， et al. The community vegetation composition and stability characteristics of alpine meadow under two grazing modes. Acta Prataculturae Sinica， 2019， 28（9）： 1-10.
	施颖，胡廷花，高红娟，等. 两种放牧模式下高寒草甸群落植被构成及稳定性特征. 草业学报， 2019， 28（9）： 1-10.
25	Lande R， Arnold S J. The measurement of selection on correlated characters. Evolutionary Ecology， 1983， 37（6）： 1210-1226.
26	Zhang B， Claßen-Bockhoff R. Sex-differential reproduction success and selection on floral traits in gynodioecious Salvia pratensis. BMC Plant Biology， 2019， 19（1）： 1-10.
27	Xie L H， Huang Q Y， Cao H J， et al. Leaf functional traits of Acer mono in Wudalianchi Volcano， China. Biodiversity Science， 2019， 27（3）： 286-296.
	谢立红，黄庆阳，曹宏杰，等. 五大连池火山色木槭叶功能性状特征. 生物多样性， 2019， 27（3）： 286-296.
28	Conner J K. Understanding natural selection： An approach integrating selection gradients， multiplicative fitness components， and path analysis. Ethology Ecology & Evolution， 1996， 8（4）： 387-397.
29	Hodgins K A， Barrett S C H. Natural selection on floral traits through male and female function in wild populations of the heterostylous daffodil Narcissus triandrus. Evolution， 2008， 62（7）： 1751-1763.
30	Gomez. Phenotypic selection and response to selection in Lobularia maritima： Importance of direct and correlational components of natural selection. Journal of Evolutionary Biology， 2001， 13（4）： 689-699.
31	Irwin R E. The consequences of direct versus indirect species interactions to selection on traits： Pollination and nectar robbing in Ipomopsis aggregata. The American Naturalist， 2006， 167（3）： 315-328.
32	Verma S K， Angadi S G， Patil V S， et al. Growth， yield and quality of chrysanthemum （Chrysanthemum morifolium Ramat.） cv. Raja as influenced by integrated nutrient management. Karnataka Journal of Agricultural Sciences. 2011， 24： 681-683.
33	Bloch D， Erhardt A. Selection toward shorter flowers by butterflies whose probosces are shorter than floral tubes. Ecology， 2008， 89（9）： 2453-2460.
34	Nilsson L A. The evolution of flowers with deep corolla tubes. Nature， 1988， 334： 147-149.
35	Johnson S， Steiner K. Long-tongued fly pollination and evolution of floral spur length in the Disa draconis complex （Orchidaceae）. Evolution， 1997， 51： 45-53.
36	Laverty T M. Bumblebee learning and flower morphology. Animal Behaviour， 1994， 47： 531-545.
37	Darwin C. On the various contrivances by which british and foreign orchids are fertilised by insects. London： John Murray， 1862： 202.
38	Hedhly A， Hormaza J I， Herrero M. The effect of temperature on pollen germination， pollen tube growth， and stigmatic receptivity in peach. Plant Biology， 2005， 7（5）： 476-483.
39	Dietrich L， Koerner C. Thermal imaging reveals massive heat accumulation in flowers across a broad spectrum of alpine taxa. Alpine Botany， 2014， 124（1）： 27-35.
40	Zhang G P， Yang M L， Cheng X X， et al. Effects of floral morphology on flower temperature increment in alpine plants. Guihaia， 2017， 37（7）： 822-828.
	张国鹏，杨明柳，程贤训，等. 高山植物花形态特征对花温度积累的影响. 广西植物， 2017， 37（7）： 822-828.
41	Yang D M， Zhang J J， Zhou D， et al. Leaf and twig functional traits of woody plants and their relationships with environmental change： A review. Chinese Journal of Ecology， 2012， 31（3）： 702-713.
	杨冬梅，章佳佳，周丹，等. 木本植物茎叶功能性状及其关系随环境变化的研究进展. 生态学杂志， 2012， 31（3）： 702-713.
42	Irwin R E. Morphological variation and female reproductive success in two sympatric Trillium species： Evidence for phenotypic selection in Trilliumerectum and Trilliumgrandiflorum （Liliaceae）. American Journal of Botany， 2000， 87（2）： 205-214.
43	Wu Y， Liu Y R， Peng H， et al. Pollination ecology of alpine herb Meconopsis integrifolia at different altitudes. Chinese Journal of Plant Ecology， 2015， 39（1）： 1-13.
	吴云，刘玉蓉，彭瀚，等. 高山植物全缘叶绿绒蒿在不同海拔地区的传粉生态学研究. 植物生态学报， 2015， 39（1）： 1-13.
44	Zhao Z G， Du G Z， Zhou X H， et al. Variations with altitude in reproductive traits and resource allocation of three tibetan species of ranunculaceae. Australian Journal of Botany， 2006， 54（7）： 691-700.
45	Milla R， Reich P B. Multi-trait interactions， not phylogeny， fine-tune leaf size reduction with increasing altitude. Annals of Botany， 2011， 107（3）： 455-465.
46	Zhang Q， Zhao C Z， Ma X L， et al. Response of reproductive allocation of Stellera chamaejasme population in alpine grassland to altitude. Chinese Journal of Ecology， 2013， 32（2）： 247-252.
	张茜，赵成章，马小丽，等. 高寒草地狼毒种群繁殖分配对海拔的响应. 生态学杂志， 2013， 32（2）： 247-252.

项目 Item	冠筒长 Corolla tube length	冠口 Corolla entrance	小花数 Flowers number	花序头直径 Inflorescence head diameter	单株丛花序数 Inflorescence number per plant
冠口 Corolla entrance	0.481**
小花数 Flowers number	0.274**	0.021
花序头直径Inflorescence head diameter	0.808**	0.511**	0.362**
单株丛花序数Inflorescence number per plant	-0.216*	-0.218*	-0.049	-0.255**
株高 Plant height	0.065	-0.050	0.234**	0.132	0.273**

项目 Item	冠筒长 Corolla tube length	冠口 Corolla entrance	小花数 Flowers number	花序头直径 Inflorescence head diameter	单株丛花序数 Inflorescence number per plant
冠口 Corolla entrance	0.481**
小花数 Flowers number	0.274**	0.021
花序头直径Inflorescence head diameter	0.808**	0.511**	0.362**
单株丛花序数Inflorescence number per plant	-0.216*	-0.218*	-0.049	-0.255**
株高 Plant height	0.065	-0.050	0.234**	0.132	0.273**

项目 Item	性状 Traits	选择梯度 Selection gradients	T值（P值） T value (P value)
定向选择Directional selection （β±SE）	冠筒长 Corolla tube length	0.067±0.040	1.669 （0.095)
	冠口 Corolla entrance	0.053±0.041	1.300 （0.194)
	单花序小花数Flowers number of per inflorescence	0.303±0.041	7.391 （0.000)
	单株丛花序数Inflorescence number of per plant	-0.007±0.048	-0.142 （0.887)
	株高 Plant height	-0.102±0.039	-2.634 （0.008)
非线性选择Nonlinear selection (2γ±SE)	冠筒长 Corolla tube length	-0.164±0.076	-2.152 （0.031)
	冠口Corolla entrance	-0.010±0.052	-0.181 （0.856)
	单花序小花数 Flowers number of per inflorescence	-0.138±0.060	-2.281 （0.023)
	单株丛花序数Inflorescence number of per plant	-0.008±0.066	-0.119 （0.905)
	株高 Plant height	0.012±0.060	0.206 （0.837)

项目 Item	性状 Traits	选择梯度 Selection gradients	T值（P值） T value (P value)
定向选择Directional selection （β±SE）	冠筒长 Corolla tube length	0.067±0.040	1.669 （0.095)
	冠口 Corolla entrance	0.053±0.041	1.300 （0.194)
	单花序小花数Flowers number of per inflorescence	0.303±0.041	7.391 （0.000)
	单株丛花序数Inflorescence number of per plant	-0.007±0.048	-0.142 （0.887)
	株高 Plant height	-0.102±0.039	-2.634 （0.008)
非线性选择Nonlinear selection (2γ±SE)	冠筒长 Corolla tube length	-0.164±0.076	-2.152 （0.031)
	冠口Corolla entrance	-0.010±0.052	-0.181 （0.856)
	单花序小花数 Flowers number of per inflorescence	-0.138±0.060	-2.281 （0.023)
	单株丛花序数Inflorescence number of per plant	-0.008±0.066	-0.119 （0.905)
	株高 Plant height	0.012±0.060	0.206 （0.837)

项目 Item	性状 Traits	选择梯度 Selection gradients	T值 (P值） T value (P value)
定向选择Directional selection (β±SE)	冠筒长 Corolla tube length	0.094±0.035	2.715 (0.006)
	单花序小花数 Flowers number of per inflorescence	0.291±0.038	7.689 (0.000)
	株高 Plant height	-0.108±0.034	-3.210 (0.001)
非线性选择Nonlinear selection (2γ±SE)	冠筒长Corolla tube length	-0.124±0.056	-2.247 (0.024)
非线性选择Nonlinear selection (2γ±SE)	单花序小花数Flowers number of per inflorescence	-0.152±0.048	-3.168 (0.001)
相关选择Correlational selection	株高×冠筒长Plant height×corolla tube length	0.059±0.037	1.618 (0.106)