Eric G. Lamb

Plants integrate information about nutrients and neighbors.

Plant root growth is modified in the presence of within-species competition and uneven local resource distributions. Animals regularly integrate information about the location of resources and the presence of competitors, altering their foraging behavior accordingly. We studied the annual plant Abutilon theophrasti to determine whether a plant can demonstrate a similarly complex response to two conditions: presence of a competitor and heterogeneous resource distributions. Individually grown plants fully explored the pot by using a broad and uniform rooting distribution regardless of soil resource distributions. Plants with competitors and uniform soil nutrient distributions exhibited pronounced reductions in rooting breadth and spatial soil segregation among the competing individuals. In contrast, plants with competitors and heterogeneous soil nutrient distributions reduced their root growth only modestly, indicating that plants integrate information about both neighbor and resource distributions in determining their root behavior.

DIRECT AND INDIRECT CONTROL OF GRASSLAND COMMUNITY STRUCTURE BY LITTER, RESOURCES, AND BIOMASS

Multiple factors linked through complex networks of interaction including fertilization, aboveground biomass, and litter control the diversity of plant communities. The challenge of explaining plant diversity is to determine not only how each individual mechanism directly influences diversity, but how those mechanisms indirectly influence diversity through interactions with other mechanisms. This approach is well established in the study of plant species richness, but surprisingly little effort has been dedicated toward understanding the controls of community evenness, despite the recognition that this aspect of diversity can influence a variety of critical ecosystem functions. Similarly, studies of diversity have predominantly focused on the influence of shoot, rather than root, biomass, despite the fact that the majority of plant biomass is belowground in many natural communities. In this study, I examine the roles of belowground biomass, live aboveground biomass, litter, and light availability in controlling the species richness and evenness of a rough fescue grassland community using structural equation modeling. Litter was the primary mechanism structuring grassland diversity, with both richness and evenness declining with increasing litter cover. There were few relationships between shoot biomass, shading, and diversity, and more importantly, no relationship between root biomass and diversity. The lack of relationship between root biomass and species richness and evenness suggests that, even though root competition in grasslands is intense, belowground interactions may not play an important role in structuring community diversity or composition.

When Competition Does Not Matter: Grassland Diversity and Community Composition

We examined whether the intense root competition in a rough fescue grassland plant community in central Alberta, Canada, was important in structuring plant species diversity or community composition. We measured competition intensity across gradients of species richness, evenness, and community composition, using pairs of naturally occurring plants of 12 species. One plant in each pair was isolated from neighbors to measure competition; community structure and environmental conditions were also measured at each pair. We used structural equation modeling to examine how competition influenced community structure. Competition intensity was unrelated to species richness and community composition, but increased competition intensity was associated with a slight decline in evenness. Size‐symmetric root competition was probably unimportant in structuring this plant community because there are no feedback mechanisms through which size‐symmetric competition can magnify small initial differences and eventually lead to competitive exclusion. In plant communities with little shoot competition, competition and community structure should be unlinked regardless of competition intensity. In more productive systems, we propose that interactions between root and shoot competition may indirectly structure communities by altering the overall asymmetry of competition.

Effects of plant species richness and evenness on soil microbial community diversity and function

Understanding the links between plant diversity and soil communities is critical to disentangling the mechanisms by which plant communities modulate ecosystem function. Experimental plant communities varying in species richness, evenness, and density were established using a response surface design and soil community properties including bacterial and archaeal abundance, richness, and evenness were measured. The potential to perform a representative soil ecosystem function, oxidation of ammonium to nitrite, was measured via archaeal and bacterial amoA genes. Structural equation modeling was used to explore the direct and indirect effects of the plant community on soil diversity and potential function. Plant communities influenced archaea and bacteria via different pathways. Species richness and evenness had significant direct effects on soil microbial community structure, but the mechanisms driving these effects did not include either root biomass or the pools of carbon and nitrogen available to the soil microbial community. Species richness had direct positive effects on archaeal amoA prevalence, but only indirect impacts on bacterial communities through modulation of plant evenness. Increased plant evenness increased bacterial abundance which in turn increased bacterial amoA abundance. These results suggest that plant community evenness may have a strong impact on some aspects of soil ecosystem function. We show that a more even plant community increased bacterial abundance, which then increased the potential for bacterial nitrification. A more even plant community also increased total dissolved nitrogen in the soil, which decreased the potential for archaeal nitrification. The role of plant evenness in structuring the soil community suggests mechanisms including complementarity in root exudate profiles or root foraging patterns.

Increased competition does not lead to increased phylogenetic overdispersion in a native grassland

That competition is stronger among closely related species and leads to phylogenetic overdispersion is a common assumption in community ecology. However, tests of this assumption are rare and field-based experiments lacking. We tested the relationship between competition, the degree of relatedness, and overdispersion among plants experimentally and using a field survey in a native grassland. Relatedness did not affect competition, nor was competition associated with phylogenetic overdispersion. Further, there was only weak evidence for increased overdispersion at spatial scales where plants are likely to compete. These results challenge traditional theory, but are consistent with recent theories regarding the mechanisms of plant competition and its potential effect on phylogenetic structure. We suggest that specific conditions related to the form of competition and trait conservatism must be met for competition to cause phylogenetic overdispersion. Consequently, overdispersion as a result of competition is likely to be rare in natural communities.

Water and nitrogen addition differentially impact plant competition in a native rough fescue grassland

We examined how water and nitrogen addition and water–nitrogen interactions affect root and shoot competition intensity and competition–productivity relationships in a native rough fescue grassland in central Alberta, Canada. Water and nitrogen were added in a factorial design to plots and root exclusion tubes and netting were used to isolate root and shoot competition on two focal species (Artemisia frigida and Chenopodium leptophyllum). Both water and nitrogen were limiting to plant growth, and focal plant survival rates increased with nitrogen but not water addition. Relative allocation to root biomass increased with water addition. Competition was almost entirely belowground, with focal plants larger when released from root but not shoot competition. There were no significant relationships between productivity and root, shoot, or total competition intensity, likely because in this system shoot biomass was too low to cause strong shoot competition and root biomass was above the levels at which root competition saturates. Water addition had few effects on the intensity of root competition suggesting that root competition intensity is invariant along soil moisture gradients. Contrary to general expectation, the strength of root competition increased with nitrogen addition demonstrating that the relationship between root competition intensity and nitrogen is more complex than a simple monotonic decline as nitrogen increases. Finally, there were few interactions between nitrogen and water affecting competition. Together these results indicate that the mechanisms of competition for water and nitrogen likely differ.

Structural equation modeling in the plant sciences: An example using yield components in oat

Lamb, E. G., Shirtliffe, S. J. and May, W. E. 2011. Structural equation modeling in the plant sciences: An example using yield components in oat. Can. J. Plant Sci. 91: 603-619. Structural equation modeling (SEM) is a powerful statistical approach for the analysis of complex intercorrelated data with a wide range of potential applications in the plant sciences. In this paper we introduce plant scientists to the principles and practice of SEM using as an example an agronomic field trial. We briefly review the history of SEM and path analysis and introduce the statistical concepts underlying SEM. We demonstrate the use of observed and latent variable structural equation models using a multi-site multi-year field trial examining the effects of seed size and seeding density on the plant density and yield of oat in Saskatchewan. Using SEM allowed for insights that a standard univariate analysis would not have revealed. We show that seeding density has strong effects on plant and panicle density, but has very limited effects on final yield. Plant density has a consistent non-linear effect on panicle density across location that was not affected by precipitation. In contrast, the implicit effect of precipitation on seed number appears to be the main driver for final yield. Incorporating precipitation data into the model demonstrates how mechanistic models can be developed by including in the path diagram variables that would normally treated as random factors in a mixed model analysis. Finally, we provide a guideline to assist plant scientists in determining whether SEM is an appropriate method to be used for the analysis of their data.

Bryophyte-cyanobacterial associations as a key factor in N2-fixation across the Canadian Arctic

Nitrogen inputs via biological N2-fixation are important in arctic environments where N often limits plant productivity. An understanding of the direct and indirect theoretical causal relationships between key intercorrelated variables that drive the process of N2-fixation is essential to understanding N input. An exploratory multi-group Structural Equation Modeling (SEM) approach was used to examine the direct and indirect effects of soil moisture, plant community functional composition, and bryophyte and lichen abundance on rates of nitrogen fixation at a low arctic ecosystem, two high arctic oases and a high arctic polar desert in the Canadian Arctic. Increasing soil moisture was strongly associated with an increasing presence of bryophytes and increasing bryophyte abundance was a major factor determining higher N2-fixation rates at all sites. Shrubs had a negative effect on bryophyte abundance at all sites with the exception of the polar desert site at Alexandra Fjord highland. The importance of competition from vascular plants appears to be greater in more productive sites and may increase at lower latitudes. Moisture availability may have an indirect effect on ecosystem development by affecting N input into the system with bryophyte-cyanobacterial associations playing an important intermediary role in the process.

Plant species traits across a riparian-zone/forest ecotone

Abstract We examined the changes in prevalence of nine plant traits – including the presence of woody stem tissue, leaf longevity, nitrogen fixation, seed longevity, dispersal vector, pollination vector, and clonal growth form – across a riparian/forest-understory ecotone. This ecotone, found along headwater streams in boreal mixed-wood forests, supports four distinct vegetation zones: streambank, riparian, transition, and upland forest understory. The objective of this study was to identify specific trait patterns that may indicate functional responses to the changes in environmental factors such as nutrient availability and wind exposure that occur across the ecotone. The suites of plant species traits found in each zone were distinct, with a strong change in the prevalence of several traits. Wind and insect pollination, wind and vertebrate diaspore dispersal, and deciduous and evergreen leaves showed the greatest change in prevalence between the vegetation types. Some traits, including insect pollination and vertebrate diaspore dispersal, were strongly correlated within species. The consistent co-occurrence of pairs of traits in the same species suggests common responses by very different traits to the same environmental factor. This study demonstrates that an ecotone can be characterized not only as a discontinuity in species distributions or environmental factors, but also as a discontinuity in the trait spectrum. Examining ecotones from a trait perspective has strong potential for identifying the environmental factors and associated species functional responses that encourage the development of distinct vegetation boundaries. Nomenclature: Newmaster et al. (1998).

Smooth brome invasion increases rare soil bacterial species prevalence, bacterial species richness and evenness

Summary Plant and soil communities are tightly linked, but the mechanisms by which the invasion of an exotic plant and the resulting shifts in plant diversity and productivity influence soil bacterial community structure remain poorly understood. We investigated the effects of invasive smooth brome (Bromus inermis) on grassland soil bacterial community structure using massively parallel sequencing of the 16S rRNA gene to determine bacterial community richness, evenness, composition and beta diversity (UniFrac indices) of soils collected along a gradient of smooth brome abundance. We evaluated several hypotheses including: (a) that the declines in native plant diversity associated with smooth brome invasion would cause declines in bacterial community diversity and (b) that mechanisms driving smooth brome effects on bacterial community structure involved altered soil edaphic properties rather than preferential invasion in areas of high soil nitrogen and distinct soil microbial communities. Smooth brome invasion led to increased soil nitrogen, soil carbon and root biomass. Bacterial evenness and bacterial richness increased with increasing smooth brome cover, while bacterial beta diversity declined. We found no evidence of a dominant direct link between the alteration of soil edaphic properties by brome and the changes in the soil bacterial community. Rather, the main controls on the soil bacterial community were direct effects of pH and smooth brome that could not be linked to the edaphic changes. The most important effect of brome on the bacterial community was the selective suppression of dominant bacterial species, which allowed rarer bacteria to increase in relative abundance. Synthesis. Here, we show that plant community composition influences bacterial community structure at a very fine scale, but that these changes are not due to altered soil total nitrogen or carbon content. The dominant direct effect of smooth brome invasion on soil communities suggests non-edaphic, that is inter- and intratrophic, interactions among smooth brome and non-bacterial components of the soil ecosystem are key drivers of soil community structure.