Predicting ecosystem changes by a new model of ecosystem evolution.

In recent years, computer simulation has been increasingly used to predict changes in actual ecosystems. In these studies, snapshots of ecosystems at certain points in time were instantly constructed without considering their evolutionary histories. However, it may not be possible to correctly predict future events unless their evolutionary processes are considered. In this study, we developed a new ecosystem model for reproducing the evolutionary process on an oceanic island, targeting Nakoudojima Island of the Ogasawara Islands. This model successfully reproduced the primitive ecosystem (the entire island covered with forest) prior to the invasion of alien species. Also, by adding multiple alien species to this ecosystem, we were able to reproduce temporal changes in the ecosystem of Nakoudojima Island after invasion of alien species. Then, we performed simulations in which feral goats were eradicated, as had actually been done on the island; these suggested that after the eradication of feral goats, forests were unlikely to be restored. In the ecosystems in which forests were not restored, arboreous plants with a high growth rate colonized during the early stage of evolution. As arboreous plants with a high growth rate consume a large amount of nutrient in soil, creating an oligotrophic state. As a result, plants cannot grow, and animal species that rely on plants cannot maintain their biomass. Consequently, many animals and plants become extinct as they cannot endure disturbances by alien species, and the ecosystem loses its resilience. Therefore, even if feral goats are eradicated, forests are not restored. Thus, the founder effect from the distant past influences future ecosystem changes. Our findings show that it is useful to consider the evolutionary process of an ecosystem in predicting its future events.

Soil phosphorous is the primary factor determining species-specific plant growth depending on soil acidity in island ecosystems with severe erosion.

Disturbances caused by invasive ungulates alter soil environments markedly and can prevent ecosystem recovery even after eradication of the ungulates. On oceanic islands, overgrazing and trampling by feral goats has caused vegetation degradation and soil erosion, which can alter soil chemistry. To understand the effects of the changes on plant performance, we conducted a laboratory experiment to assess herbaceous species growth under various soil conditions with phosphorous, nutrients, and acidity. Subsoil was collected from Nakodo-jima in the northwest Pacific. Six herbaceous species dominating the island were grown in soils with three levels of added CaCO3 and P2O5 and two levels of added KNO3. After 4 weeks of growth, the total dry plant weight was significantly lower with no added P2O5, regardless of the addition of KNO3. Three species weighed more under P2O5 and KNO3 addition in high-pH soil, whereas the remaining three weighed less. Our results indicated that herbaceous species growth is limited primarily by phosphorous availability; the limitation is dependent on soil pH, and the trend of dependency differs among species. This implies that ecosystems with extreme disturbances cannot recover without improving the soil chemistry.

Cardiac Dysfunction Due to Thiamine Deficiency after Hemodialysis for Biguanide-related Lactic Acidosis.

Biguanide is an ideal drug for the treatment of type 2 diabetes mellitus. When used appropriately, the incidence of lactic acidosis is reported to be very low. Risk factors associated with biguanide-related lactic acidosis include chronic kidney disease, congestive heart failure, alcohol use, severe dehydration, shock, hypoxic states, sepsis, and advanced age. We herein report a case of cardiac dysfunction due to thiamine deficiency after hemodialysis in a patient with suspected biguanide-related lactic acidosis. Patients who develop severe lactic acidosis while taking biguanides should be given a large dose of thiamine without delay, given the possibility of thiamine deficiency as a complication.

Functional differences in seasonally absorbed nitrogen in a winter-green perennial herb.

Nitrogen (N) uptake in response to its availability and effective N-use are important for determining plant fitness, as N is a major limiting resource and its availability changes both seasonally and annually. Storage organs such as bulbs are considered an adaptive trait with respect to plant N-use strategies. It is well known that N is remobilized from storage organs to satisfy the high demand for new growth that is not completely satisfied by external uptake alone. However, little is known about how this N absorbed during different seasons contributes to plant performance. By manipulating seasonal N availability in potted Lycoris radiata var. radiata (Amaryllidaceae), a winter-green perennial, we found that the N absorbed during different seasons had different effects on leaf growth and leaf N concentrations, effectively increasing the growth and survival of the plants. N absorbed during the summer (leafless period; N was thus stored in the bulb) enhanced plant growth by increasing leaf growth. Compared with the plants supplied with N during autumn (leaf flush period), the leafy plants also showed greater growth per unit leaf area despite the lower area-based photosynthetic capacity of the latter. By contrast, N absorbed during the autumn increased the leaf N concentration and thus the photosynthetic capacity, which was considered to enhance survival and growth of the plant during winter by reducing the potentially fatal risk caused by the absorption of photons under low temperature. Our findings have important implications for estimating plant responses to environmental changes. We predict that changes in seasonal N availability impact the performance of plants, even that of perennials that have large storage organs, via an altered relative investment of N into different functions.

Distance- and density-dependent leaf dynamics of seedlings of a tropical rainforest tree.

Parental distance and plant density dependence of seedling leaf turnover and survival was examined to investigate predictions of the Janzen-Connell hypothesis. The focal study species, Shorea macroptera is a canopy tree species in a lowland rain forest in peninsular Malaysia. We found that the peak of the distribution of plants shifted from 3-6 m to 6-9 m during the course of the change from seedling to sapling stage. The leaf demography of the seedlings was influenced by their distance from the adult tree and also by the seedling density. Although significant density- and distance dependence in leaf production was not detected, seedling leaf loss decreased with distance from the parent tree and with seedling density. Similarly, leaf damage was not found to be distance- or density-dependent, but net leaf gain of seedlings increased with distance from the parent tree. Although no significant distance- or density-dependence was evident in terms of leaf damage, significant distance dependence of the net leaf gain was found. Thus, we concluded that positive distance dependence in the leaf turnover of seedlings may gradually contribute to a shift in the distribution pattern of the progeny through reductions in growth and survivorship.

Increases in soil water content after the mortality of non-native trees in oceanic island forest ecosystems are due to reduced water loss during dry periods.

The control of dominant, non-native trees can alter the water balance of soils in forest ecosystems via hydrological processes, which results in changes in soil water environments. To test this idea, we evaluated the effects of the mortality of an invasive tree, Casuarina equisetifolia Forst., on the water content of surface soils on the Ogasawara Islands, subtropical islands in the northwestern Pacific Ocean, using a manipulative herbicide experiment. Temporal changes in volumetric water content of surface soils at 6 cm depth at sites where all trees of C. equisetifolia were killed by herbicide were compared with those of adjacent control sites before and after their mortality with consideration of the amount of precipitation. In addition, the rate of decrease in the soil water content during dry periods and the rate of increase in the soil water content during rainfall periods were compared between herbicide and control sites. Soil water content at sites treated with herbicide was significantly higher after treatment than soil water content at control sites during the same period. Differences between initial and minimum values of soil water content at the herbicide sites during the drying events were significantly lower than the corresponding differences in the control quadrats. During rainfall periods, both initial and maximum values of soil water contents in the herbicided quadrats were higher, and differences between the maximum and initial values did not differ between the herbicided and control quadrats. Our results indicated that the mortality of non-native trees from forest ecosystems increased water content of surface soils, due primarily to a slower rate of decrease in soil water content during dry periods.

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.

Growth strategies differentiate the spatial patterns of 11 dipterocarp species coexisting in a Malaysian tropical rain forest.

We examined relationships between mortality rate, relative growth rate (RGR), and spatial patterns of three growth stages (small, medium, and large trees) for 11 dipterocarp species in the Pasoh 50-ha plot. Mortality rates for these species tended to be positively correlated with RGRs, although the correlation was significant only at the small-tree stage. Seven species with high growth and mortality rates exhibited peaks in spatial aggregation at small distances (<100 m) in small trees, but this aggregation disappeared in medium and large trees. In contrast, the other four species with low growth and mortality rates aggregated at large distances (>200 m) throughout the three growth stages in all but one species. Negative associations between different growth stages were observed only for the high-mortality species, suggesting density-dependent mortality. The high-mortality species showed habitat associations with topography, soil type, and the forest regeneration phase after gap formation, whereas the three low-mortality species only had associations with the forest regeneration phase. A randomization procedure revealed that these habitat associations explained little of their spatial aggregation. Our results suggest that the growth strategy has a large effect on the structuring of the spatial distribution of tree species through mortality processes.

Variation in flowering size and age of a facultative biennial, Aster kantoensis (Compositae), in response to nutrient availability.

Although the flowering of facultative biennials is size-dependent, flowering size varies markedly within a single population as well as among populations. In this study, 15 half-sib families of the facultative biennial Aster kantoensis were grown from seeds at three nutrient levels (low, medium, and high). A significant nutrient × family interaction effect was found for bolting size, and among-family variation in bolting size increased with decreasing nutrient level. Growth from bolting to flowering tended to be greatest at the high nutrient level. Such responses of bolting size and growth from bolting to flowering resulted in an increase in flowering size at the high nutrient level and a significant variation in its reaction norm among families. For flowering age, there was a significant interaction of nutrient × family, and its among-family variation increased with decreasing nutrient levels, as was the case with bolting size. These results indicate that genetic variation in phenotypic plasticity of bolting size with nutrient availability was one cause of the variation in flowering size and age in the A. kantoensis population on the floodplain with the spatially heterogeneous nutrient availability. Moreover, responses of growth from bolting to flowering to nutrient availability could enhance the variation in flowering size.

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.

Hydrochory as a determinant of genetic distribution of seeds within Hibiscus moscheutos (Malvaceae) populations.

Seed dispersal is a major determinant of the spatial genetic structure of plant populations. In this study, we evaluated the role of distinct hydrologic regimes in determining the spatial genetic structure of the seed bank of the wetland plant Hibiscus moscheutos. We analyzed seeds in surface soil samples collected in the autumn and the following spring by determining their allozyme genotypes and estimated the pattern in seed movements during flooding. We selected study sites in nontidal and tidal wetlands with different flooding regimes. One nontidal site had no flooding, while the second nontidal site was inundated for most of the year. One tidal wetland site flooded with almost every tide, and a second tidal site was inundated at moderate frequency. Genetic makeup of the seed bank at the nonflooded site changed little between seasons. Secondary seed dispersal altered absolute allele frequencies at the other three sites, with the greatest change occurring at the two tidally influenced sites. This study demonstrates that secondary hydrochory influences the genetic composition of the seed bank and that hydrologic conditions play an important role in determining the local patterns in seed movements.

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.

Delayed greening, leaf expansion, and damage to sympatric Shorea species in a lowland rain forest.

The function of delayed greening in the seedlings of canopy tree species in a lowland tropical rain forest was examined in terms of its potential defensive value against herbivory. To explore the ecological and evolutionary backgrounds for delayed greening, we chose eight sympatric congeneric (Shorea) dipterocarp species that were either normal-greening or delayed-greening species. Expansion and toughening of leaves took approximately 30 days for all species, and did not differ between the normal- and delayed-greening species. The main factors that affected leaf damage during expansion were insect herbivory and fungal infection. Levels of leaf damage were significantly lower for delayed-greening species than for normal-greening species, but proportions of heavily damaged leaves and leaf abscission during expansion did not differ. In addition, no significant difference was found in damage levels on leaves (aged 1-2 months) of naturally occurring seedlings between normal- and delayed-greening species. Therefore, delayed greening may effectively reduce the level of leaf damage in young expanding leaves, but may not necessarily reduce leaf abscission and damage to mature leaves. The existence of delayed greening could not be simply explained by the phylogenetic and ecological backgrounds of the trees. Consequently, delayed greening may have a function in reducing damage during expansion, but more information (such as knowledge of the secondary metabolites involved in this phenomenon) is needed to explain fully why these species exhibit delayed greening.

Temporal and spatial patterns of mass flowerings on the Malay Peninsula.

We propose a hypothesis to explain the temporal and spatial patterns of mass flowerings in dipterocarp tree species on the Malay Peninsula. The literature on these mass flowerings reveals that during 1980-2002 at least 11 flowerings occurred at irregular intervals of 1-6 yr in a lowland rain forest. Five of them were typical mass flowerings-a high density of flowering trees and the characteristic sequential flowering of Shorea species. The 11 flowerings were classified into two flowering times: spring and autumn. There is evidence that low temperature and drought triggered the flowerings. Therefore, the seasonality of mass flowerings is characterized by the annual patterns of rainfall and low temperature. In addition, changes in El Niño-Southern Oscillation (ENSO) may play important roles in determining the supra-annual occurrence of mass flowerings. Flowering surveys on the Malay Peninsula implied that regions with spring or autumn mass flowerings corresponded geographically to those regions that had one cool season (December-February) or two (December-February and June-August), respectively. This finding anticipates the seasonal pattern and geographical distribution of mass flowerings on the Malay Peninsula.