A review on the relationships between plant genetic diversity and ecosystem functioning

doi:10.17521/cjpe.2018.0013

Abstract

Abstract:

The loss of genetic diversity is accelerating due to habitat loss and population reduction caused by global change and anthropologenic activities. For species-poor ecosystems, the effect of genetic diversity on ecosystem functioning may not be smaller than that of species diversity. Therefore, understanding the relationship between genetic diversity and ecosystem functioning (GD-EF) and its underlying mechanisms is important for biodiversity conservation, responses of ecosystems to environmental change and ecological restoration. Here, we reviewed the studies on the effects of plant genetic diversity on ecosystem structures (community structure of the higher tropic level) and ecosystem functions (primary production, nutrient cycling and ecosystem stability), and the mechanisms underlying these relationships. We also discussed the influence of functional diversity on GD-EF, the comparison of effects of the genetic and species diversity on ecosystem functioning, and the application of GD-EF in the ecological restorations. We finally pointed out the limitations in current studies to provide references for the future: (1) further studies on the mechanisms of GD-EF are needed; (2) no study has evaluated the influence of genetic diversity on maltifunctinarity; (3) the impacts of different measurements of genetic diversity on ecosystem functioning are unclear; (4) there are lack of long-time GD-EF studies and GD-EF studies conducted at multidimensional scales; (5) the relative importance of genetic diversity and other factors on ecosystem functioning in the nature is unclear.

Key words: genetic diversity, genotypic diversity, ecosystem structure, ecosystem function, additive effect, complementarity effect

ZHANG Li-Wen, HAN Guang-Xuan. A review on the relationships between plant genetic diversity and ecosystem functioning[J]. Chin J Plan Ecolo, 2018, 42(10): 977-989.

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URL: https://www.plant-ecology.com/EN/10.17521/cjpe.2018.0013

https://www.plant-ecology.com/EN/Y2018/V42/I10/977

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References 82

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名词术语 Glossary	解释 Explanation
遗传多样性 Genetic diversity	种群内个体间遗传变异的程度。 The degree of genetic variation among individuals in a population.
基因丰富度 Allelic richness	所检测位点上等位基因的平均数目。 The average number of alleles detected at the detected locus.
基因多样性 Allelic diversity	包含位点上基因数目和频率信息的一类遗传多样性指数, 比如: Shannon信息指数和期望杂合度。 A class of genetic diversity indices containing information about the number and frequency of genes at a locus, such as: Shannon information index and expected heterozygosity.
基因型 Genotype	一个个体在指定数量的位点上等位基因的组成。 The composition of alleles of an individual at a specified number of loci.
基因型丰富度 Genotypic richness	一个种群中基因型的数目。 The number of genotypes in a population.
基因型均匀度 Genotypic eveness	基因型多度的分布。如果一个种群中各基因型多度等同, 那么基因型均匀度为1; 如果一个种群只有一个基因型, 那么该种群基因型均匀度为0。 The distribution of genotypic abundance. The genotype evenness is 1 if genotypic abundance is equal in a population, and 0 if there is only one genotype in a population.
基因型相异度 Genotypic dissimilarity	一个种群中两两基因型间遗传距离的平均值。 The average genetic distance between two genotypes in a population.
基因型亲缘度 Genotypic relatedness	与基因型相异度相反, 指一个种群中两两基因型间亲缘程度的平均值。对于植物微卫星分子标记数据, 二倍体可以用STORM软件(Frasier, 2008)或者R程序包“related” (Pew et al., 2015), 多倍体用P_OLYR_ELATEDNESS计算基因型亲缘度(Huang et al., 2015)。 The average value of relatedness between two genotypes in a population, which is contrary to genotypic dissimilarity. For plant microsatellite marker data, genotypic relatedness of diploids can be calculated with STORM software (Frasier, 2008) or R package “related” (Pew et al., 2015), and genotypic relatedness of polyploids can be calculated with P_OLYR_ELATEDNESS (Huang et al., 2015).
适应性遗传多样性 Adaptive genetic diversity	适应性遗传多样性是通过改变表达蛋白质的数量或结构或表达时间来影响表型以帮助个体适应环境或者提高个体适合度的变异。一般采用已知遗传关系的个体(比如, 来自同一母株种子生长的个体)开展同质园数量遗传实验进行估算, 但是这个方法比较费时费力费钱; 另外一种方法是开发和筛选出适应性分子标记来测定其遗传变异。 Adaptive genetic diversity is the variation that affects phenotypes by altering the number or structure of expressed proteins or the expression time to help individuals adapt to the environment or improve their fitness. Quantitative genetic common garden experiments are usually conducted to estimate the adaptive gentic variation by using individuals with known genetic relationships (e.g., individuals from seeds of the same mother tree), but this method is time-consuming and costly; another method is to develop and select adaptive molecular markers to determine their genetic variation.
中性遗传多样性 Neutral genetic diversity	中性遗传多样性是由不影响表型的序列变异组成。中性遗传多样性和适应性遗传多样性也可能有相关性, 原因是采用的中性分子标记位点与适应性遗传变异可能存在连锁不平衡的情况。 Neutral genetic diversity is composed of sequence variations that do not affect phenotypes. Neutral genetic diversity and adaptive genetic diversity may also be correlated, because there may be linkage imbalance between the neutral molecular marker loci and adaptive genetic variation.
品种 Cultivar	为特定的某一性状或若干性状的组合而选择出来的植物集合体, 在这些性状上是特异、一致、稳定的, 并且通过适当的有性或无性方式繁殖时仍保持这些性状。 Plants were selected for a particular trait or combination of several specific traits, and these traits are specific, consistent, and stable, and retained when propagated sexually or asexually.
近交衰退 Inbreeding depression	由于近交(自交和亲缘个体间的异交)导致后代适合度下降的现象, 产生的主要原因是由于近交增加了有害等位基因的纯合几率。 Inbreeding (selfing and outcrossing between related individuals) results in a decrease in fitness of offspring, mainly because inbreeding increases the probability of harmful homozygous alleles.
远交衰退 Outbreeding depression	不同生境的种群个体, 各自拥有适应当地生境的特有等位基因组合, 如果它们相互之间杂交(交配)将可能打破这种组合, 引起后代适应能力降低。 Individuals from different habitats have specific allele combinations adapted to local habitats. If they cross breeding (mate) with each other, they may break the specific allele combinations and reduce the adaptability of their offspring.
功能多样性 Functional diversity	植物个体水平上的形态、生理以及生活史特征等功能性状通过影响植物存活能力、生长和繁殖来影响其适合度, 这些功能性状特征值的大小、范围和分布状况称为功能多样性。 Functional traits such as morphology, physiology and life history at the individual level affect plant fitness by affecting its survival, growth and reproduction. The size, range and distribution of these functional trait values are called functional diversity.
奠基者多样性 Founder diversity	这里指的是, 在种群保护或者生态修复中, 所引物种种群的遗传多样性。奠基者效应是指由少数个体的基因频率决定了它们后代基因频率的效应。 This refers to the genetic diversity of founder in population conservation or ecological restoration. Founder effect is the gene frequency of a small number of individuals determines the gene frequency of their offspring.
系统发育多样性 Phylogenetic diversity	群落中物种的系统发育树形图(表示物种之间的亲缘关系)中所有分枝长度之和。 The sum of all the branch lengths in a phylogenetic tree of species in a community (representing the relatedness between species).

名词术语 Glossary	解释 Explanation
遗传多样性 Genetic diversity	种群内个体间遗传变异的程度。 The degree of genetic variation among individuals in a population.
基因丰富度 Allelic richness	所检测位点上等位基因的平均数目。 The average number of alleles detected at the detected locus.
基因多样性 Allelic diversity	包含位点上基因数目和频率信息的一类遗传多样性指数, 比如: Shannon信息指数和期望杂合度。 A class of genetic diversity indices containing information about the number and frequency of genes at a locus, such as: Shannon information index and expected heterozygosity.
基因型 Genotype	一个个体在指定数量的位点上等位基因的组成。 The composition of alleles of an individual at a specified number of loci.
基因型丰富度 Genotypic richness	一个种群中基因型的数目。 The number of genotypes in a population.
基因型均匀度 Genotypic eveness	基因型多度的分布。如果一个种群中各基因型多度等同, 那么基因型均匀度为1; 如果一个种群只有一个基因型, 那么该种群基因型均匀度为0。 The distribution of genotypic abundance. The genotype evenness is 1 if genotypic abundance is equal in a population, and 0 if there is only one genotype in a population.
基因型相异度 Genotypic dissimilarity	一个种群中两两基因型间遗传距离的平均值。 The average genetic distance between two genotypes in a population.
基因型亲缘度 Genotypic relatedness	与基因型相异度相反, 指一个种群中两两基因型间亲缘程度的平均值。对于植物微卫星分子标记数据, 二倍体可以用STORM软件(Frasier, 2008)或者R程序包“related” (Pew et al., 2015), 多倍体用P_OLYR_ELATEDNESS计算基因型亲缘度(Huang et al., 2015)。 The average value of relatedness between two genotypes in a population, which is contrary to genotypic dissimilarity. For plant microsatellite marker data, genotypic relatedness of diploids can be calculated with STORM software (Frasier, 2008) or R package “related” (Pew et al., 2015), and genotypic relatedness of polyploids can be calculated with P_OLYR_ELATEDNESS (Huang et al., 2015).
适应性遗传多样性 Adaptive genetic diversity	适应性遗传多样性是通过改变表达蛋白质的数量或结构或表达时间来影响表型以帮助个体适应环境或者提高个体适合度的变异。一般采用已知遗传关系的个体(比如, 来自同一母株种子生长的个体)开展同质园数量遗传实验进行估算, 但是这个方法比较费时费力费钱; 另外一种方法是开发和筛选出适应性分子标记来测定其遗传变异。 Adaptive genetic diversity is the variation that affects phenotypes by altering the number or structure of expressed proteins or the expression time to help individuals adapt to the environment or improve their fitness. Quantitative genetic common garden experiments are usually conducted to estimate the adaptive gentic variation by using individuals with known genetic relationships (e.g., individuals from seeds of the same mother tree), but this method is time-consuming and costly; another method is to develop and select adaptive molecular markers to determine their genetic variation.
中性遗传多样性 Neutral genetic diversity	中性遗传多样性是由不影响表型的序列变异组成。中性遗传多样性和适应性遗传多样性也可能有相关性, 原因是采用的中性分子标记位点与适应性遗传变异可能存在连锁不平衡的情况。 Neutral genetic diversity is composed of sequence variations that do not affect phenotypes. Neutral genetic diversity and adaptive genetic diversity may also be correlated, because there may be linkage imbalance between the neutral molecular marker loci and adaptive genetic variation.
品种 Cultivar	为特定的某一性状或若干性状的组合而选择出来的植物集合体, 在这些性状上是特异、一致、稳定的, 并且通过适当的有性或无性方式繁殖时仍保持这些性状。 Plants were selected for a particular trait or combination of several specific traits, and these traits are specific, consistent, and stable, and retained when propagated sexually or asexually.
近交衰退 Inbreeding depression	由于近交(自交和亲缘个体间的异交)导致后代适合度下降的现象, 产生的主要原因是由于近交增加了有害等位基因的纯合几率。 Inbreeding (selfing and outcrossing between related individuals) results in a decrease in fitness of offspring, mainly because inbreeding increases the probability of harmful homozygous alleles.
远交衰退 Outbreeding depression	不同生境的种群个体, 各自拥有适应当地生境的特有等位基因组合, 如果它们相互之间杂交(交配)将可能打破这种组合, 引起后代适应能力降低。 Individuals from different habitats have specific allele combinations adapted to local habitats. If they cross breeding (mate) with each other, they may break the specific allele combinations and reduce the adaptability of their offspring.
功能多样性 Functional diversity	植物个体水平上的形态、生理以及生活史特征等功能性状通过影响植物存活能力、生长和繁殖来影响其适合度, 这些功能性状特征值的大小、范围和分布状况称为功能多样性。 Functional traits such as morphology, physiology and life history at the individual level affect plant fitness by affecting its survival, growth and reproduction. The size, range and distribution of these functional trait values are called functional diversity.
奠基者多样性 Founder diversity	这里指的是, 在种群保护或者生态修复中, 所引物种种群的遗传多样性。奠基者效应是指由少数个体的基因频率决定了它们后代基因频率的效应。 This refers to the genetic diversity of founder in population conservation or ecological restoration. Founder effect is the gene frequency of a small number of individuals determines the gene frequency of their offspring.
系统发育多样性 Phylogenetic diversity	群落中物种的系统发育树形图(表示物种之间的亲缘关系)中所有分枝长度之和。 The sum of all the branch lengths in a phylogenetic tree of species in a community (representing the relatedness between species).

效应类别(按生物学角度来分类) Types of genetic diversity effects (in terms of biology)	效应区别 The differences among types of genetic diversity effect	生态系统功能 Ecosystem functioning	相关机制假说 Hypothesis	假说内容 Content of hypothesis
加性效应 Additive effects	不同基因型对生态系统功能的效应是独立的、可加的; 基因型间相互作用不影响其生态系统功能效应。 The effects of different genotypes on ecosystem functioning are independent and additive, and the interactions among genotypes do not affect their effects on ecosystem functioning.	高营养级生物群落结构、初级生产力、养分循环、生态系统稳定性 Community structure of higher trophic levels, primary productivity, nutrientcycling and ecosystem stability	优势效应 Dominant effect	基因型多样性高的系统中包含对系统有利的基因型概率高于单基因型系统, 因此基因型多样性高有利于维持生态系统结构和功能。验证方法请见1.1和1.2节。 Systems with high genotypic diversity have a higher probability of containing genotypes beneficial to the system than those with single genotype, and thus high genotypic diversity is conducive to maintaining ecosystem structure and function. Please see sections 1.1 and 1.2 for the methods for testing this hypotheses.
非加性效应 Non-additive effects	不同基因型对生态系统功能效应是非独立的、不可加的; 基因型间相互作用对生态系统功能产生交互效应。The effects of different genotypes on ecosystem functioning are dependent and nonadditive, and the interactions among genotypes affect their effects on ecosystem functioning.	高营养级生物群落结构 Community structure of higher trophic levels	资源化假说 The resource specialization hypothesis	大多数植食性节肢动物表现出一定程度的食性偏好, 随着基因型多样性的增加, 相关联的植食性节肢动物多样性也增加。验证方法请见1.1节。 Most of the herbivorous arthropods showed food preference. With the increase of genotypic diversity, the diversity of associated herbivorous arthropods also increased. Please see Section 1.1 for the methods for testing this hypothesis.
		高营养级生物群落结构 Community structure of higher trophic levels	更多个体假说 The more individuals hypothesis	如果地上净生产力随着基因型多样性升高, 那么能够提供更多的能量给更多植食性节肢动物, 这些植食性节肢动物多样性随之增加, 继而捕食者也会增加。验证方法请见1.1节。 If net aboveground productivity increases with genotypic diversity, more energy can be provided to more herbivorous arthropods, and the diversity of these herbivorous arthropods increases, followed by increased predators. Please see Section 1.1 for the methods for testing this hypothesis.
		初级生产力、养分循环、生态系统稳定性 Primary productivity, nutrient cycling and ecosystem stability	性状独立互补效应 Trait-indepen-dent complementarity	在多基因型系统, 如果某些基因型的生态系统功能比它们在单基因型系统提高了, 但与基因型的功能性状无关, 而且不以抑制其他基因型的适合度为代价(不同基因型占据不重叠的生态位形成生态位互补或者基因型间存在正作用), 则为正效应。验证方法请见1.2节。 In genotypic mixture, if the ecosystem functions of some genotypes are improved than that in the genotypic monoculture but are not related to the functional traits of the genotypes, and are also not at the expense of inhibiting the fitness of other genotypes (different genotypes occupy non-overlapping niches to form niche complementarities or have positive effects between genotypes). It is a positive effect. Please see section 1.2 for the methods for testing this hypothesis.
			性状依赖互补效应 Trait-dependent complementarity	在多基因型系统, 如果具有特殊功能性状基因型(比如, 特殊功能性状使得基因型间形成嵌套生态位)的生态功能比其在单基因型系统增加了, 且不以抑制其他基因型适合度为代价, 则为正效应。验证方法请见1.2节。 In genotypic mixture, if the ecological functions of genotypes with special functional traits (for example, the nested niches formed between genotypes) are increased than those of genotypic monoculture, and not at the expense of inhibiting the fitness of other genotypes, the effect is positive. Please see section 1.2 the methods for testing this hypothesis.

效应类别(按生物学角度来分类) Types of genetic diversity effects (in terms of biology)	效应区别 The differences among types of genetic diversity effect	生态系统功能 Ecosystem functioning	相关机制假说 Hypothesis	假说内容 Content of hypothesis
加性效应 Additive effects	不同基因型对生态系统功能的效应是独立的、可加的; 基因型间相互作用不影响其生态系统功能效应。 The effects of different genotypes on ecosystem functioning are independent and additive, and the interactions among genotypes do not affect their effects on ecosystem functioning.	高营养级生物群落结构、初级生产力、养分循环、生态系统稳定性 Community structure of higher trophic levels, primary productivity, nutrientcycling and ecosystem stability	优势效应 Dominant effect	基因型多样性高的系统中包含对系统有利的基因型概率高于单基因型系统, 因此基因型多样性高有利于维持生态系统结构和功能。验证方法请见1.1和1.2节。 Systems with high genotypic diversity have a higher probability of containing genotypes beneficial to the system than those with single genotype, and thus high genotypic diversity is conducive to maintaining ecosystem structure and function. Please see sections 1.1 and 1.2 for the methods for testing this hypotheses.
非加性效应 Non-additive effects	不同基因型对生态系统功能效应是非独立的、不可加的; 基因型间相互作用对生态系统功能产生交互效应。The effects of different genotypes on ecosystem functioning are dependent and nonadditive, and the interactions among genotypes affect their effects on ecosystem functioning.	高营养级生物群落结构 Community structure of higher trophic levels	资源化假说 The resource specialization hypothesis	大多数植食性节肢动物表现出一定程度的食性偏好, 随着基因型多样性的增加, 相关联的植食性节肢动物多样性也增加。验证方法请见1.1节。 Most of the herbivorous arthropods showed food preference. With the increase of genotypic diversity, the diversity of associated herbivorous arthropods also increased. Please see Section 1.1 for the methods for testing this hypothesis.
		高营养级生物群落结构 Community structure of higher trophic levels	更多个体假说 The more individuals hypothesis	如果地上净生产力随着基因型多样性升高, 那么能够提供更多的能量给更多植食性节肢动物, 这些植食性节肢动物多样性随之增加, 继而捕食者也会增加。验证方法请见1.1节。 If net aboveground productivity increases with genotypic diversity, more energy can be provided to more herbivorous arthropods, and the diversity of these herbivorous arthropods increases, followed by increased predators. Please see Section 1.1 for the methods for testing this hypothesis.
		初级生产力、养分循环、生态系统稳定性 Primary productivity, nutrient cycling and ecosystem stability	性状独立互补效应 Trait-indepen-dent complementarity	在多基因型系统, 如果某些基因型的生态系统功能比它们在单基因型系统提高了, 但与基因型的功能性状无关, 而且不以抑制其他基因型的适合度为代价(不同基因型占据不重叠的生态位形成生态位互补或者基因型间存在正作用), 则为正效应。验证方法请见1.2节。 In genotypic mixture, if the ecosystem functions of some genotypes are improved than that in the genotypic monoculture but are not related to the functional traits of the genotypes, and are also not at the expense of inhibiting the fitness of other genotypes (different genotypes occupy non-overlapping niches to form niche complementarities or have positive effects between genotypes). It is a positive effect. Please see section 1.2 for the methods for testing this hypothesis.
			性状依赖互补效应 Trait-dependent complementarity	在多基因型系统, 如果具有特殊功能性状基因型(比如, 特殊功能性状使得基因型间形成嵌套生态位)的生态功能比其在单基因型系统增加了, 且不以抑制其他基因型适合度为代价, 则为正效应。验证方法请见1.2节。 In genotypic mixture, if the ecological functions of genotypes with special functional traits (for example, the nested niches formed between genotypes) are increased than those of genotypic monoculture, and not at the expense of inhibiting the fitness of other genotypes, the effect is positive. Please see section 1.2 the methods for testing this hypothesis.

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