克隆和有性亲体效应及其调控机制

植物表型受自身基因型、所处环境及其亲体所经历环境的共同影响；其中，亲体环境对子代表型的影响被称为亲体效应。亲体效应不仅可通过有性繁殖产生的种子传递给后代（即有性亲体效应），也可以通过克隆生长等无性繁殖产生的分株传递给后代（即克隆亲体效应）。亲体效应对植物种群，特别是对有性繁殖受限、缺乏遗传变异的克隆植物种群的长期进化可能发挥着极其重要的作用，因此，对亲体效应研究进展的梳理非常必要。对克隆亲体效应和有性亲体效应的内涵进行了阐释，并论述了克隆和有性亲体效应对子代表型、适合度、种内/种间竞争能力以及种群/群落结构和功能的潜在影响；阐述了亲体效应的潜在调控机制，包括供给机制、代谢物质调控机制、表观遗传机制等；论述了克隆亲体效应在克隆植物适应进化中的作用。未来可以就克隆亲体效应的遗传稳定性及其对克隆生活史性状变异的贡献程度，以及克隆和有性亲体效应引起的表型多样性对种内/种间关系、种群/群落多样性及生态系统结构、功能和稳定性的影响开展深入研究。

Clonal and sexual parental effects and their mechanisms

The phenotype of a plant can be jointly determined by its genotype, the environmental condition experienced by itself and the environmental condition(s) experienced by its parents. The latter, i.e., the impacts of parental environment conditions on offspring phenotypes, are referred to as parental effects, maternal effects or paternal effects. Because parental effects reflect the response of offspring phenotypes to environment conditions across generations, it is also named transgenerational plasticity. Parental effects may have played an important role in the long-term evolution of plant populations, especially for those with limited sexual reproduction and thus low genetic variation. In this paper, we reviewed the research progresses of parental effects and pointed out the future research directions. We explicitly defined two types of parental effects, i.e., clonal parental effects and sexual parental effects. The former refer to parental effects that are transmitted to offspring through ramets produced by clonal growth and other asexual reproduction, while the latter refer to those that are transmitted to offspring through seeds produced by sexual reproduction. While sexual parental effects have long been recognized, clonal parental effects are called for attention only in recent years. Both sexual and clonal parental effects generally enhance offspring performance, but sometimes also show neutral or even negative effects, in response to abiotic or biotic stresses. For clonal parental effects, a few studies have investigated that the impacts on growth and physiological traits of clonal offspring strongly depend on offspring environment and levels of plants (i.e., an individual offspring or all offspring of a parent); however, such impacts on exclusive life-history traits of clonal offspring, such as clonal growth, clonal integration, clonal plasticity and intraclonal labor division, are still unclear. The potential mechanisms of parental effects are the provisioning mechanism, metabolic regulation mechanism and epigenetic mechanism. In natural conditions, these three mechanisms often jointly act to affect offspring phenotypes, and their effects cannot be strictly separated from each other. Although a large body of studies have examined parental effects at individual level, there are still limited studies at population level and virtually none at community level. As parental effects can influence offspring phenotypes, they can influence their intra- and inter-specific competitive ability, which may further impact population structure and dynamics and community function and stability. Therefore, future studies of parental effects should test the impact of phenotypic differences/diversity induced by parental effects on intra-/inter-specific relationships, population structure and dynamics and community function (e.g., productivity) and stability. The epigenetic mechanism underlying these population- and community-level parental effects should also be explicitly evaluated.