青藏高原东缘龙胆属植物繁殖分配对策研究
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摘要
资源分配的模式可以反映植物生活史特征,是植物生活史研究的核心,近年来,对资源分配的研究主要集中在性分配和繁殖分配两方面。性分配主要是研究投入到繁殖中的资源如何在雌雄功能之间分配的问题;繁殖分配主要研究植物体总资源中供给繁殖部分的比例、影响这种比例的因素以及繁殖分配与这些影响因素之间的关系。
青藏高原东部高寒草甸的特殊地理环境及气候条件,孕育了许多独特的植物类群。这些植物如何适应青藏高原的胁迫环境,表现出什么样的繁殖对策,引起了繁殖生态学家和进化生物学家的极大兴趣。本文通过对广泛分布于该区域的龙胆属植物常见种的资源分配,尤其是繁殖对策与各种影响因素关系的研究,试图揭示该属植物对环境的适应对策。结果表明:
(1)龙胆属植物的植株个体越大,繁殖投入越高,花大小和总花数也随个体增大而增加,其中花数量的增加比花大小的增加更加显著,而繁殖分配与个体大小间无固定的相关模式,表明不同物种对相似环境压力的适应对策不同,对于多年生龙胆来说,随着植物个体的增大,对雌性、雄性和吸引结构的投入都在增加,这可保证资源的充分利用,不会因为单一部分的增加而造成资源的浪费;
(2) 6种多年生龙胆属植物中,有4种其性分配结果与SDS的理论预测一致,即大个体更偏向雌性器官的资源投入,但麻花艽(Gentiana atraminea)和达乌里秦艽(Gentiana dahurica)的性分配与个体大小则没有表现出负相关,可能与其本身具有的雌雄异熟——雄性先熟特点有关;
(3)资源在雌雄功能间的分配没有表现出权衡关系,可能是由于植物必须在许多不同生活史性状之间进行资源分配,而不是两两之间非此即彼。在采用偏相关方法控制植物个体大小后,花大小与花数目之间表现出权衡关系,与理论预测一致。
(4)除反折花龙胆的个体大小随海拔升高而降低外,所研究种的个体大小与繁殖投入均随海拔升高表现出正相关趋势,说明龙胆属植物倾向于采取增加个体大小与繁殖投入的方式来应对随海拔升高所造成的环境胁迫的增强。海拔高度对繁殖分配的影响因种而异,可见对于高海拔环境,龙胆属植物采取了不同的繁殖分配对策。
(5)本研究中的龙胆属植物,其P/O比均随海拔升高而显著降低,高海拔环境传粉者的缺乏很可能是造成这种繁育系统对策变化的主要选择压力,此外,除麻花艽外,随着海拔升高龙胆属植物的花数量变化不大,花的大小则有增大趋势,说明在花大小与数量所代表的雌雄繁殖适合度方面,植物偏向于雄性适合度的增加。
(6)从原始种到进化种它们的繁殖分配比例表现出逐渐上升的趋势,这表明进化程度越高的种,对于繁殖的相对投入越高;而一年生龙胆较多年生龙胆的繁殖分配比例显著提高说明一年生植物更加重视繁殖方面的输出,以求在有限的生活史中,获得尽可能大的适合度。
(7)在P/O值方面,多年生龙胆的P/O高于一年生龙胆,单看多年生龙胆的几个种,除秦艽组两个种外,随着进化程度的提高,P/O逐渐增加,一年生龙胆也表现出相似的结果,这说明龙胆属植物的P/O同样与其系统发育水平有联系,而秦艽组的例外则可能与所研究两个种正好都是聚伞花序有关。
The mode of resources allocation can reflect the character of the plant life history, which is central to study of plant life history. Studies on resource allocation were focused on two aspects: sex allocation and reproductive allocation. Sex allocation is about optimal allocation of reproductive resources between the male and female; Reproductive allocation is about the ratio of reproductive resources in total, the factors which influence the ratio, and the relationship between the ratio and the influence factors.
There are a lot of unique plants in Alpine Meadow at Eastern Qinghai Tibetan Plateau. How to adapt the especial environment of Alpine, and what reproductive strategies appear for these species, which raised great interest of breeding ecologist and evolutionary biologist. Therefore, we selected common species of Gentiana, which has widely distribution at eastern of Qinghai-Tibet Plateau, studied the resource allocation strategies, especially the correlation between reproductive strategies and various influencing factors, to understand how the plants adapt to the especial environmental condition. The results showed that:
(1) The larger of individual size, the higher of reproductive investment and both floral display size and total flower production are increasing with plant size , the flower number is increasing more remarkably than flower size. But there was not certain correlation between reproductive allocation and size of individuals, which suggest species-specific adaptive strategies. For perennial plants of Gentiana, with increasing of individual size, female function investment, male function investment and attractive structures investment increase. It can ensure that all the resource will be available fully without waste due to the increasing of a single part;
(2)There are four results of sex allocation in the six kinds of perennial gentiana concordant with SDS theory prediction, that is to say, the larger individual allocates more resources to female function. The sex allocations of Gentiana atraminea and Gentiana dahurica Show no negative correlation with the individual size may be related to the Dichogamy-Protandry;
(3)The distribution between female and male function of resource show no trade-off. It maybe caused by the truth that the resource distribution of plant must be under diverse life history character, not one kind with another. The trade-off between total flower production and flower size is exist after controlling for plant size statistically. This result confirms the theory predicts;
(4) The positively correlated between individuals size、reproductive investment and altitudes, except plant size of Gentiana Choanantha. It means that plants of Gentiana tend to increase the plant size and reproductive investment to adapt the varied environment. Influences induced by high altitude on the reproductive allocation vary by species, which suggest species-specific adaptive strategies.
(5) In this work, with the altitude increased the pollen/ovule ratio of perennial gentiana reduced, and the shortages of pollinators at a high altitude may be the main pressure for forming this breeding system countermeasure changes. Moreover, except Gentiana atraminea, the variety of the amount of flowers has little influence on the altitude increased, but the size of flowers is trending up. This shows that the plant of gentian usually favored male fitness on male and female reproduction fitness which is represented by flower size and flower number.
(6)The proportion of reproduction allocation of gentiana shows a tendency to increasing gradually from original species to evolution species. It means that the higher development of species, the more devotion for reproduction. Comparing with perennial gentiana, the proportion of reproduction allocation of annual gentiana has risen notably. It means that the annual plants usually pay more attention on the reproduction output to gain maximum fitness during its limited life cycle.
(7) For the pollen/ovule ratio, the perennial gentiana is higher than annual gentiana. Excepting the two species of Sect. Cruciata, the pollen/ovule ratio of other species of perennial gentiana increased by higher development. The annual gentiana also has the same feature. It means that the pollen/ovule ratio of gentiana also associated with its phylogeny level. The exception of Sect. Cruciata maybe related on that the two species which are studied by us are cyme.
青藏高原东部高寒草甸的特殊地理环境及气候条件,孕育了许多独特的植物类群。这些植物如何适应青藏高原的胁迫环境,表现出什么样的繁殖对策,引起了繁殖生态学家和进化生物学家的极大兴趣。本文通过对广泛分布于该区域的龙胆属植物常见种的资源分配,尤其是繁殖对策与各种影响因素关系的研究,试图揭示该属植物对环境的适应对策。结果表明:
(1)龙胆属植物的植株个体越大,繁殖投入越高,花大小和总花数也随个体增大而增加,其中花数量的增加比花大小的增加更加显著,而繁殖分配与个体大小间无固定的相关模式,表明不同物种对相似环境压力的适应对策不同,对于多年生龙胆来说,随着植物个体的增大,对雌性、雄性和吸引结构的投入都在增加,这可保证资源的充分利用,不会因为单一部分的增加而造成资源的浪费;
(2) 6种多年生龙胆属植物中,有4种其性分配结果与SDS的理论预测一致,即大个体更偏向雌性器官的资源投入,但麻花艽(Gentiana atraminea)和达乌里秦艽(Gentiana dahurica)的性分配与个体大小则没有表现出负相关,可能与其本身具有的雌雄异熟——雄性先熟特点有关;
(3)资源在雌雄功能间的分配没有表现出权衡关系,可能是由于植物必须在许多不同生活史性状之间进行资源分配,而不是两两之间非此即彼。在采用偏相关方法控制植物个体大小后,花大小与花数目之间表现出权衡关系,与理论预测一致。
(4)除反折花龙胆的个体大小随海拔升高而降低外,所研究种的个体大小与繁殖投入均随海拔升高表现出正相关趋势,说明龙胆属植物倾向于采取增加个体大小与繁殖投入的方式来应对随海拔升高所造成的环境胁迫的增强。海拔高度对繁殖分配的影响因种而异,可见对于高海拔环境,龙胆属植物采取了不同的繁殖分配对策。
(5)本研究中的龙胆属植物,其P/O比均随海拔升高而显著降低,高海拔环境传粉者的缺乏很可能是造成这种繁育系统对策变化的主要选择压力,此外,除麻花艽外,随着海拔升高龙胆属植物的花数量变化不大,花的大小则有增大趋势,说明在花大小与数量所代表的雌雄繁殖适合度方面,植物偏向于雄性适合度的增加。
(6)从原始种到进化种它们的繁殖分配比例表现出逐渐上升的趋势,这表明进化程度越高的种,对于繁殖的相对投入越高;而一年生龙胆较多年生龙胆的繁殖分配比例显著提高说明一年生植物更加重视繁殖方面的输出,以求在有限的生活史中,获得尽可能大的适合度。
(7)在P/O值方面,多年生龙胆的P/O高于一年生龙胆,单看多年生龙胆的几个种,除秦艽组两个种外,随着进化程度的提高,P/O逐渐增加,一年生龙胆也表现出相似的结果,这说明龙胆属植物的P/O同样与其系统发育水平有联系,而秦艽组的例外则可能与所研究两个种正好都是聚伞花序有关。
The mode of resources allocation can reflect the character of the plant life history, which is central to study of plant life history. Studies on resource allocation were focused on two aspects: sex allocation and reproductive allocation. Sex allocation is about optimal allocation of reproductive resources between the male and female; Reproductive allocation is about the ratio of reproductive resources in total, the factors which influence the ratio, and the relationship between the ratio and the influence factors.
There are a lot of unique plants in Alpine Meadow at Eastern Qinghai Tibetan Plateau. How to adapt the especial environment of Alpine, and what reproductive strategies appear for these species, which raised great interest of breeding ecologist and evolutionary biologist. Therefore, we selected common species of Gentiana, which has widely distribution at eastern of Qinghai-Tibet Plateau, studied the resource allocation strategies, especially the correlation between reproductive strategies and various influencing factors, to understand how the plants adapt to the especial environmental condition. The results showed that:
(1) The larger of individual size, the higher of reproductive investment and both floral display size and total flower production are increasing with plant size , the flower number is increasing more remarkably than flower size. But there was not certain correlation between reproductive allocation and size of individuals, which suggest species-specific adaptive strategies. For perennial plants of Gentiana, with increasing of individual size, female function investment, male function investment and attractive structures investment increase. It can ensure that all the resource will be available fully without waste due to the increasing of a single part;
(2)There are four results of sex allocation in the six kinds of perennial gentiana concordant with SDS theory prediction, that is to say, the larger individual allocates more resources to female function. The sex allocations of Gentiana atraminea and Gentiana dahurica Show no negative correlation with the individual size may be related to the Dichogamy-Protandry;
(3)The distribution between female and male function of resource show no trade-off. It maybe caused by the truth that the resource distribution of plant must be under diverse life history character, not one kind with another. The trade-off between total flower production and flower size is exist after controlling for plant size statistically. This result confirms the theory predicts;
(4) The positively correlated between individuals size、reproductive investment and altitudes, except plant size of Gentiana Choanantha. It means that plants of Gentiana tend to increase the plant size and reproductive investment to adapt the varied environment. Influences induced by high altitude on the reproductive allocation vary by species, which suggest species-specific adaptive strategies.
(5) In this work, with the altitude increased the pollen/ovule ratio of perennial gentiana reduced, and the shortages of pollinators at a high altitude may be the main pressure for forming this breeding system countermeasure changes. Moreover, except Gentiana atraminea, the variety of the amount of flowers has little influence on the altitude increased, but the size of flowers is trending up. This shows that the plant of gentian usually favored male fitness on male and female reproduction fitness which is represented by flower size and flower number.
(6)The proportion of reproduction allocation of gentiana shows a tendency to increasing gradually from original species to evolution species. It means that the higher development of species, the more devotion for reproduction. Comparing with perennial gentiana, the proportion of reproduction allocation of annual gentiana has risen notably. It means that the annual plants usually pay more attention on the reproduction output to gain maximum fitness during its limited life cycle.
(7) For the pollen/ovule ratio, the perennial gentiana is higher than annual gentiana. Excepting the two species of Sect. Cruciata, the pollen/ovule ratio of other species of perennial gentiana increased by higher development. The annual gentiana also has the same feature. It means that the pollen/ovule ratio of gentiana also associated with its phylogeny level. The exception of Sect. Cruciata maybe related on that the two species which are studied by us are cyme.
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