Can heritable traits in a single species affect an entire ecosystem? Recent studies show that such traits in a common tree have predictable effects on community structure and ecosystem processes. Because these 'community and ecosystem phenotypes' have a genetic basis and are heritable, we can begin to apply the principles of population and quantitative genetics to place the study of complex communities and ecosystems within an evolutionary framework. This framework could allow us to understand, for the first time, the genetic basis of ecosystem processes, and the effect of such phenomena as climate change and introduced transgenic organisms on entire communities.
Mycorrhizal fungi influence plant growth, local biodiversity and ecosystem function. Effects of the symbiosis on plants span the continuum from mutualism to parasitism. We sought to understand this variation in symbiotic function using meta-analysis with information theory-based model selection to assess the relative importance of factors in five categories: (1) identity of the host plant and its functional characteristics, (2) identity and type of mycorrhizal fungi (arbuscular mycorrhizal vs. ectomycorrhizal), (3) soil fertility, (4) biotic complexity of the soil and (5) experimental location (laboratory vs. field). Across most subsets of the data, host plant functional group and N-fertilization were surprisingly much more important in predicting plant responses to mycorrhizal inoculation (Ôplant responseÕ) than other factors. Non-N-fixing forbs and woody plants and C 4 grasses responded more positively to mycorrhizal inoculation than plants with N-fixing bacterial symbionts and C 3 grasses. In laboratory studies of the arbuscular mycorrhizal symbiosis, plant response was more positive when the soil community was more complex. Univariate analyses supported the hypothesis that plant response is most positive when plants are P-limited rather than N-limited. These results emphasize that mycorrhizal function depends on both abiotic and biotic context, and have implications for plant community theory and restoration ecology.
We present evidence that the heritable genetic variation within individual species, especially dominant and keystone species, has community and ecosystem consequences. These consequences represent extended phenotypes, i.e., the effects of genes at levels higher than the population. Using diverse examples from microbes to vertebrates, we demonstrate that the extended phenotype can be traced from the individuals possessing the trait, to the community, and to ecosystem processes such as leaf litter decomposition and N mineralization. In our development of a community genetics perspective, we focus on intraspecific genetic variation because the extended phenotypes of these genes can be passed from one generation to the next, which provides a mechanism for heritability. In support of this view, common‐garden experiments using synthetic crosses of a dominant tree show that their progeny tend to support arthropod communities that resemble those of their parents. We also argue that the combined interactions of extended phenotypes contribute to the among‐community variance in the traits of individuals within communities. The genetic factors underlying this among‐community variance in trait expression, particularly those involving genetic interactions among species, constitute community heritability. These findings have diverse implications. (1) They provide a genetic framework for understanding community structure and ecosystem processes. The effects of extended phenotypes at these higher levels need not be diffuse; they may be direct or may act in relatively few steps, which enhances our ability to detect and predict their effects. (2) From a conservation perspective, we introduce the concept of the minimum viable interacting population (MVIP), which represents the size of a population needed to maintain genetic diversity at levels required by other interacting species in the community. (3) Genotype × environment interactions in dominant and keystone species can shift extended phenotypes to have unexpected consequences at community and ecosystem levels, an issue that is especially important as it relates to global change. (4) Documenting community heritability justifies a community genetics perspective and is an essential first step in demonstrating community evolution. (5) Community genetics requires and promotes an integrative approach, from genes to ecosystems, that is necessary for the marriage of ecology and genetics. Few studies span from genes to ecosystems, but such integration is probably essential for understanding the natural world. Corresponding Editor: A. A. Agrawal
Summary1 Although drought frequency and severity are predicted to increase across numerous continental interiors, the consequences of these changes for dominant plants are largely unknown. Over the last decade, the south-western US has experienced six drought years, including the extreme droughts of 1996 and 2002, which led to widespread tree mortality across northern Arizona. 2 We examined the impact of these droughts on the co-dominant tree species of the pinyon-juniper woodland ( Pinus edulis and Juniperus monosperma ), a major vegetation type in the US. 3 Pinyon mortality following both droughts was 6.5-fold higher than juniper mortality. In addition, large pinyons suffered 2-6-fold greater mortality than small pinyons, a pattern associated with higher mortality of reproductively mature trees and survival of smaller pinyons resulting from facilitation by established vegetation. Differential mortality of large pinyons resulted in a vegetation shift such that the pinyon-juniper woodlands are becoming dominated by juniper, a species that is typical of lower elevations and more arid conditions. 4 Sites that experienced high pinyon mortality during the first drought suffered additional mortality during the second drought, so that reductions in tree densities and the resulting release from below-ground competition did not buffer surviving pinyons against additional mortality during the second drought. Such repeated mortality events also suggest that these stands may suffer chronic stress. 5 Reductions in biotic associations (e.g. avian seed dispersers, ectomycorrhizas and nurse plants) that will probably result from extreme mortality of large pinyons ensure that the observed vegetation shifts will be persistent. Because approximately 1000 species are associated with pinyon pine, the shift in the structure of these woodlands has large-scale community consequences.
Studies that combine experimental manipulations with long-term data collection reveal elaborate interactions among species that affect the structure and dynamics of ecosystems. Research programs in U.S. desert shrubland and pinyon-juniper woodland have shown that (i) complex dynamics of species populations reflect interactions with other organisms and fluctuating climate; (ii) genotype x environment interactions affect responses of species to environmental change; (iii) herbivore-resistance traits of dominant plant species and impacts of "keystone" animal species cascade through the system to affect many organisms and ecosystem processes; and (iv) some environmental perturbations can cause wholesale reorganization of ecosystems because they exceed the ecological tolerances of dominant or keystone species, whereas other changes may be buffered because of the compensatory dynamics of complementary species.
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