Resource allocation in tragedy of the commons game in plants for belowground competition.

The tragedy of the commons (TOC) has been well known since it was proposed and has been widely applied not only to human society but also to many taxa. An increasing number of studies have focused on TOC in belowground competition in plants. In the presence of neighbors, plants overproduce roots to acquire more nutrients than their competitors, resulting in a reduction in reproductive yield. Game-theoretic studies on TOC in plants usually consider the amount of root biomass as a strategy and do not consider the growth of plants. However, root volume is considered an outcome of the decision-making of plants on whether they allocate more resources to the root. In this study, we incorporated resource allocation and growth dynamics into the TOC game in plants and explored the evolutionarily stable resource allocation strategy in the presence of neighbors. We demonstrated that TOC generally occurs when fitness per individual is always reduced because of the competitive response. However, the overproliferation of roots, which is emphasized as an indicator of TOC, did not necessarily occur, or was sometimes difficult to detect when fitness is largely or completely determined by root biomass. This result suggests the importance of careful observation for examining whether plant species engage in a TOC game.

Female and male fitness consequences of clonal growth in a dwarf bamboo population with a high degree of clonal intermingling.

BACKGROUND AND AIMS: Although many studies have reported that clonal growth interferes with sexual reproduction as a result of geitonogamous self-pollination and inbreeding depression, the mating costs of clonal growth are expected to be reduced when genets are spatially intermingled with others. This study examined how clonal growth affects both female and male reproductive success by studying a population of a mass-flowering plant, Sasa veitchii var. hirsuta, with a high degree of clonal intermingling. METHODS: In a 10 × 10 m plot, genets were discriminated based on the multilocus genotypes of 11 nuclear microsatellite loci. The relationships between genet size and the components of reproductive success were then investigated. Male siring success and female and male selfing rates were assessed using paternity analysis. KEY RESULTS: A total of 111 genets were spatially well intermingled with others. In contrast to previous studies with species forming distinct monoclonal patches, seed production linearly increased with genet size. While male siring success was a decelerating function of genet size, selfing rates were relatively low and not related to genet size. CONCLUSIONS: The results, in conjunction with previous studies, emphasize the role of the spatial arrangement of genets on both the quantity and quality of offpsring, and suggest that an intermingled distribution of genets can reduce the mating costs of clonal growth and enhance overall fitness, particularly female fitness.

Clonal structure, seed set, and self-pollination rate in mass-flowering bamboo species during off-year flowering events.

Bamboos are typical examples of highly synchronized semelparous species. Their mass-flowering events occur at supra-annual intervals but they sometimes flower on a small scale in off-years. If some bamboo ramets (culms) of a genet flower and die in off-years, whereas other culms of the same genet do not flower synchronously, the genet can still survive blooming in an off-year and could participate in the next mass-flowering event. At genet level, the effect might be similar to that achieved by synchronously reproducing iteroparous plants. In addition, if multiple genets flower simultaneously in off-years, cross-pollination will be promoted. However, it is not known whether all the culms in a genet flower synchronously and whether multiple genets flower in off-years. We determined the clonal structure of three temperate dwarf bamboo species, i.e., Sasa senanensis, S. kurilensis, and S. palmata, at 24 off-year flowering sites and the surrounding areas in northern Japan using seven microsatellite markers. We also estimated seed set at seven of the sites and self-pollination rates at five sites to determine off-year reproductive success. Next, we investigated whether seed sets at the culm level were related to flowering area and/or number of flowering genets, using generalized linear mixed-effect models (GLMMs). Multiple genets flowered at 9/24 flowering sites. We found that 40/96 of the genets identified had some flowering culms. Non-flowering culms were present in 24/40 flowering genets. Seed set was in the range 2.2%-12.5% and the self-pollination rate was 96.3%. In the best GLMM, seed set increased with flowering area. Seeds were produced in off-years, but cross-pollination was rare in off-years. We suggest that some dwarf bamboos may exhibit iteroparity or imperfectly synchronized semelparity at the genet level, a characteristic similar to that of other reproductively synchronous plants. We also found synchronous flowering of a few genets even in off-years.

Availability and temporal heterogeneity of water supply affect the vertical distribution and mortality of a belowground herbivore and consequently plant growth.

We examined how the volume and temporal heterogeneity of water supply changed the vertical distribution and mortality of a belowground herbivore, and consequently affected plant biomass. Plantago lanceolata (Plantaginaceae) seedlings were grown at one per pot under different combinations of water volume (large or small volume) and heterogeneity (homogeneous water conditions, watered every day; heterogeneous conditions, watered every 4 days) in the presence or absence of a larva of the belowground herbivorous insect, Anomala cuprea (Coleoptera: Scarabaeidae). The larva was confined in different vertical distributions to top feeding zone (top treatment), middle feeding zone (middle treatment), or bottom feeding zone (bottom treatment); alternatively no larva was introduced (control treatment) or larval movement was not confined (free treatment). Three-way interaction between water volume, heterogeneity, and the herbivore significantly affected plant biomass. With a large water volume, plant biomass was lower in free treatment than in control treatment regardless of heterogeneity. Plant biomass in free treatment was as low as in top treatment. With a small water volume and in free treatment, plant biomass was low (similar to that under top treatment) under homogeneous water conditions but high under heterogeneous ones (similar to that under middle or bottom treatment). Therefore, there was little effect of belowground herbivory on plant growth under heterogeneous water conditions. In other watering regimes, herbivores would be distributed in the shallow soil and reduced root biomass. Herbivore mortality was high with homogeneous application of a large volume or heterogeneous application of a small water volume. Under the large water volume, plant biomass was high in pots in which the herbivore had died. Thus, the combinations of water volume and heterogeneity affected plant growth via the change of a belowground herbivore.

Combined effects between temporal heterogeneity of water supply, nutrient level, and population density on biomass of four broadly distributed herbaceous species.

Temporal heterogeneity of water supply affects grassland community productivity and it can interact with nutrient level and intraspecific competition. To understand community responses, the responses of individual species to water heterogeneity must be evaluated while considering the interactions of this heterogeneity with nutrient levels and population density. We compared responses of four herbaceous species grown in monocultures to various combinations of water heterogeneity, nutrient level, and population density: two grasses (Cynodon dactylon and Lolium perenne), a forb (Artemisia princeps), and a legume (Trifolium repens). Treatment effects on shoot and root biomass were analyzed. In all four species, shoot biomass was larger under homogeneous than under heterogeneous water supply. Shoot responses of L. perenne tended to be greater at high nutrient levels. Although root biomass was also larger under homogeneous water supply, effects of water heterogeneity on root biomass were not significant in the grasses. Trifolium repens showed marked root responses, particularly at high population density. Although greater shoot and root growth under homogeneous water supply appears to be a general trend among herbaceous species, our results suggested differences among species could be found in the degree of response to water heterogeneity and its interactions with nutrient level and intraspecific competition.

Effects of temporal heterogeneity of water supply on the growth of Perilla frutescens depend on plant density.

BACKGROUND AND AIMS: Plants respond to the spatial and temporal heterogeneity of a resource supply. However, their responses will depend on intraspecific competition for resource acquisition. Although plants are subject to various intensities of intraspecific competition, most studies of resource heterogeneity have been carried out under a single density so that the effects of intraspecific competition on plant responses to resource heterogeneity are largely unknown. METHODS: A growth experiment was performed to investigate plant responses to the temporal heterogeneity of water supply and nutrient levels under multiple plant densities. The annual plant Perilla frutescens was grown using different combinations of frequency of water supply, nutrient level and density, while providing the same total amount of water under all conditions. The effects of the treatments on biomass, allocation to roots and intensity of competition were analysed after 48 d. KEY RESULTS: Biomass and allocation to roots were larger under homogeneous than under heterogeneous water supply, and the effects of water heterogeneity were greater at high density than at low density. The effects of water heterogeneity were greater at high nutrient level than at low level for biomass, while the effects were greater at low nutrient level than high level for allocation to roots. Competition was severer under homogeneous than under heterogeneous water supply. CONCLUSIONS: Competition for water probably makes plants more sensitive to the water heterogeneity. In addition, the intensity of intraspecific competition can be affected by the temporal patterns of water supply. Because both resource heterogeneity and intraspecific competition affect resource acquisition and growth of plants, their interactive effects should be evaluated more carefully under future studies.

Root growth and plant biomass in Lolium perenne exploring a nutrient-rich patch in soil.

We investigated soil exploration by roots and plant growth in a heterogeneous environment to determine whether roots can selectively explore a nutrient-rich patch, and how nutrient heterogeneity affects biomass allocation and total biomass before a patch is reached. Lolium perenne L. plants were grown in a factorial experiment with combinations of fertilization (heterogeneous and homogeneous) and day of harvest (14, 28, 42, or 56 days after transplanting). The plant in the heterogeneous treatment was smaller in its mean total biomass, and allocated more biomass to roots. The distributions of root length and root biomass in the heterogeneous treatment did not favor the nutrient-rich patch, and did not correspond to the patchy distribution of inorganic nitrogen. Specific root length (length/biomass) was higher and root elongation was more extensive both laterally and vertically in the heterogeneous treatment. These characteristics may enable plants to acquire nutrients efficiently and increase the probability of encountering nutrient-rich patches in a heterogeneous soil. However, heterogeneity of soil nutrients would hold back plant growth before a patch was reached. Therefore, although no significant selective root placement in the nutrient-rich patch was observed, plant growth before reaching nutrient-rich patches differed between heterogeneous and homogeneous environments.

Development of a local size hierarchy causes regular spacing of trees in an even-aged Abies forest: analyses using spatial autocorrelation and the mark correlation function.

BACKGROUND AND AIMS: During the development of an even-aged plant population, the spatial distribution of individuals often changes from a clumped pattern to a random or regular one. The development of local size hierarchies in an Abies forest was analysed for a period of 47 years following a large disturbance in 1959. METHODS: In 1980 all trees in an 8 x 8 m plot were mapped and their height growth after the disturbance was estimated. Their mortality and growth were then recorded at 1- to 4-year intervals between 1980 and 2006. Spatial distribution patterns of trees were analysed by the pair correlation function. Spatial correlations between tree heights were analysed with a spatial autocorrelation function and the mark correlation function. The mark correlation function was able to detect a local size hierarchy that could not be detected by the spatial autocorrelation function alone. KEY RESULTS: The small-scale spatial distribution pattern of trees changed from clumped to slightly regular during the 47 years. Mortality occurred in a density-dependent manner, which resulted in regular spacing between trees after 1980. The spatial autocorrelation and mark correlation functions revealed the existence of tree patches consisting of large trees at the initial stage. Development of a local size hierarchy was detected within the first decade after the disturbance, although the spatial autocorrelation was not negative. Local size hierarchies that developed persisted until 2006, and the spatial autocorrelation became negative at later stages (after about 40 years). CONCLUSIONS: This is the first study to detect local size hierarchies as a prelude to regular spacing using the mark correlation function. The results confirm that use of the mark correlation function together with the spatial autocorrelation function is an effective tool to analyse the development of a local size hierarchy of trees in a forest.

Change in spatial distribution patterns of a biennial plant between growth stages and generations in a patchy habitat.

BACKGROUND AND AIMS: The aim of the study was to evaluate factors causing change in spatial distribution patterns of plants between growth stages and generations for a monocarpic biennial plant, Lysimachia rubida. It was assumed that habitat heterogeneity was a primary factor determining spatial patterns of plants, and a randomization procedure was developed for testing the null hypothesis that only spatial association with ground surface conditions determined spatial patterns of plants. METHODS: A 5-year demographic census was conducted on an open dry habitat that was heterogeneous with regard to the ground surface conditions. KEY RESULTS: There was significant habitat association in that plants at vegetative and reproductive stages were denser in areas with smaller gravel than with larger gravel. Point process analyses rejected the null hypothesis of the spatial association with ground surface conditions. CONCLUSIONS: The results suggest that other factors, such as patchy seed dispersal, secondary dispersal of the seeds and life-history variation at various spatial scales, also affected spatial patterns of individuals in a population of L. rubida. Spatial structures and dynamics of a local population in a patchy habitat represent various performances of plants within patches and seed dispersal within a patch and beyond the patch.

Latitudinal variation in plant size and relative growth rate in Arabidopsis thaliana.

Latitude is an important determinant of local environmental conditions that affect plant growth. Forty ecotypes of Arabidopsis thaliana were selected from a wide range of latitudes (from 16°N to 63°N) to investigate genetic variation in plant size and relative growth rate (RGR) along a latitudinal gradient. Plants were grown in a greenhouse for 31 days, during which period three consecutive harvests were performed. Plants from high latitudes tended to have smaller plant size in terms of seed size, cotyledon width, rosette size, number of rosette leaves, size (leaf area) of the largest leaves, total leaf area, and total dry weight per plant than those from low latitudes. The mean (±SE) RGR across ecotypes was 0.229 (±0.0013) day-1. There was, however, significant ecotypic variation, with RGR being negatively correlated with latitude. The two main components of RGR, leaf area ratio (LAR) and unit leaf rate (ULR), were also correlated with latitude: LAR increased with increasing latitude while ULR decreased with increasing latitude. It was also found that RGR tended to be negatively correlated with LAR, specific leaf area (SLA) and specific root length (SRL) but to be positively correlated with mean area per leaf (MAL) and ULR. The variation in RGR among ecotypes was relatively small compared with that in the other traits. RGR may be a conservative trait, whose variation is constrained by the trade-off between its physiological (i.e. ULR) and morphological (i.e. LAR) components.