Timing of disturbance changes the balance between growth and survival of parent and offspring ramets in the clonal forest understory herb Uvularia perfoliata

Preformation of organs involves the initiation of vegetative and generative tissues at least one season before they are actually produced. It is a strategy to deal with environments characterized by predictable seasonality as it enables fast growth of plants at the onset of favorable conditions. However, early preformation also strongly restricts plants in their response to unpredictable environmental changes and disturbance. In this study we investigated the response of the clonal forest understory herb Uvularia perfoliata to disturbance and resource limitation. In U. perfoliata shoot characteristics, as well as vegetative and sexual reproduction are determined at the end of the previous growing season. Plants were grown under two light levels and the rhizome connection between parent and offspring ramets were severed at various times during the growing period. Disturbance did not affect total biomass accumulation but it did affect the relative allocation and survival probability of parents and offspring ramets. Early severing resulted in increased survival chance and future fitness of the parent ramet, while late severing resulted in a higher survival chance and increased fitness of offspring ramets. The response was mediated by plant size and resource availability. These results show that the life history of U. perfoliata includes the possibility to alleviate the effects of disturbance even though the species is characterized by strong developmental canalization through organ preformation.Co-ordinating editor: J. Tuomi     

Resource sharing among ramets in the clonal herb,Fragaria chiloensis

Summary The herbaceous perennial, Fragaria chiloensis, reproduces vegetatively on coastal sand dunes in California by growth of stolons that bear rosettes. Movement of water and photosynthates through stolons integrates water and carbon metabolism of rosettes both before and after they root. New, unrooted rosettes import sufficient water and nitrogen to maintain levels near those of established rosettes; yet support of an unrooted rosette did not decrease growth of a connected, rooted sibling given abundant light, water, and soil nutrients. Under such conditions strings of unrooted rosettes with the associated stolon appeared self-sufficient for carbon; shade and drought induced import of photosynthates. New rosettes produced and maintained a limited root mass upon contact with dry sand, which could increase probability of establishment. Rooting did not induce senescence of stolons. Connection between two established rosettes prevented death by drought and shade, even when neither rosette could have survived singly. Results suggest that physiological integration of connected rosettes may increase total growth of clones of F. chiloensis through sharing of resources among ramets, especially when resource availability is changeable or patchy.     

Effect of canopy gap light environment on evaporative load and stomatal conductance in the temperate forest understory herb Aster macrophyllus (Asteraceae)

Aster macrophyllus, a temperate forest understory species of the northeastern United States, inhabits a broad range of light habitats. Plants receiving several minutes of direct sun in canopy gap and forest edge habitats occasionally wilt, a response indicative of water stress. We compared two alterative scenarios for patterns of evaporative load and stomatal conductance for plants in large (0.15 ha) tree canopy gaps and small (3 m²) herbaceous subcanopy gaps: 1) evaporative loads are typically moderate and stomatal conductance is largely governed by light intensity; or 2) evaporative loads are often substantial, mandating stomatal closure to prevent excessive transpiration. In all cases evaporative loads were elevated by light intensity above 25% of full sun. This was accompanied by substantial stomatal closure. Transitions from low to moderate light intensity (<13% full sun) caused little increase in leaf evaporative load, and stimulated increases in stomatal conductance. Very brief periods of high light also stimulated stomatal opening. Light environments in the small herbaceous subcanopy gaps differ greatly in their patterns of evaporative load from day to day.     

Antioxidant activity of compounds from the medicinal herb Aster tataricus

A number of compounds were isolated from the medicinal plant Aster tataricus including shionone, epifriedelinol, quercetin, kaempferol, scopoletin, emodin, aurantiamide acetate and 1,7-dihydroxy-6-methyl-anthraquinone. The compounds were compared with regard to their ability in inhibiting hemolysis of rat erythrocytes induced by 2'-2' azobis (2-amidinopropane) dihydrochloride, lipid peroxidation using the FeSO(4)-ascorbic acid system, and generation of superoxide radicals using a phenazine methosulfate-nicotinamide adenine dinucleotide system. The effects on the Fe-bleomycin-induced DNA damage reflected pro-oxidant activity. Quercetin and kaempferol were most potent in inhibiting hemolysis, lipid peroxidation and superoxide radical generation. Scopoletin and emodin were similar to quercetin and kaempferol in inhibiting superoxide radical generation and second to them in inhibiting lipid peroxidation. Aurantiamide acetate exhibited some inhibitory activity toward superoxide radical generation. 1,7-dihydroxy-6-methyl-anthraquinone exerted an inhibitory activity only on superoxide radical generation. Shionone and epifriedelinol did not display any antioxidant activity. Quercetin and kaempferol, but not the remaining compounds, exhibited some pro-oxidant activity.     

Fission and the dynamics of genets and ramets in clonal cnidarian populations

Until relatively recently many species of entomostracan crustaceans were thought to have widespread, even cosmopolitan distributions. Evidence now suggests that this is far less the case than thought. However, an exception appears to be provided by the crustaceans of episodically filled waterbodies. Typically these include Anostraca, Notostraca and Conchostraca. The paper considers the distribution of Triops australiensis (Branchiopoda: Notostraca) in Australia. There, it is a frequent inhabitant of episodically filled temporary waters. It is absent from such waters in northern Australia. A biogeoclimatic analysis using the program BIOCLIM indicates that this is because of the absence of suitable localities. BIOCLIM enables the preparation of maps which provide statistical predictions of climates suitable for the survival of a particular taxon. These predictions are based upon the known climate of localities where the taxon in question actually occurs. It is suggested that for the biota of shallow ephemeral waters (such as those typically inhabited by Anostraca, Conchostraca and Notostraca) biogeoclimatic analysis has a powerful predictive value.     

Intraspecific variation in the population dynamics and growth of the limpet,Cellana tramoserica

Summary Populations of the limpet Cellana tramoserica (Sowerby) from high, mid, and low intertidal regions, and from a subtidal zone, were studied at Cape Banks (N.S.W.), Australia. Individuals from the subtidal population had the largest mean and maximum shell size, the low and high shore populations were intermediate, while the midshore population had both the smallest mean and maximum size. The density of adults showed the reverse trend: the midshore region had the greatest adult population density while the subtidal population had the smallest density. The density of juveniles and recruits was negatively correlated with tidal height in the intertidal areas, but the density of both age-classes was smallest in the subtidal region. The rate of growth of individuals was negatively correlated to the adult density of a site; individuals from the subtidal population grew the fastest, while those from the midshore region grew the slowest. The level of adult mortality of the four populations was similar, with an annual rate of between 50–60%. Juvenile mortality did, however, differ among populations; reduced percentages of juveniles reached adult size in the mid and low shore populations than in the highshore and subtidal populations.The expected lifetime fecundity of individual females differed among the four populations. Subtidal individuals were expected to spawn 40 times the amount of gonad material during their life than were individuals from the midshore population. Individuals from the high and low populations probably spawn 10 times more than those from the midshore region. Differences in the fecundities of individuals were not likely to be the result of genetic differences because experimental transplants and manipulations of density showed that individuals from all the intertidal populations could increase their growth rate to match that of subtidal individuals. The implications of the difference in lifetime fecundity among populations are discussed.     

An analysis of the costs and benefits of physiological integration between ramets in the clonal perennial herb Glechoma hederacea

Summary The costs and benefits, measured in terms of dry weight, of physiological integration between clonal ramets, were analysed in two experiments conducted on the clonal herb Glechoma hederacea. Firstly, integration between consecutively-produced ramets was examined in an experiment in which stolons grew from one set of growing conditions (either unshaded or shaded and either nutrient-rich or nutrient-poor) into conditions in which light or nutrient level was altered. Comparisons were made between the dry weight of the parts of the clones produced before and after growing conditions were changed, and the dry weights of the corresponding part of control clones subjected to constant growing conditions. In a second experiment, integration between two distinct parts of G. hederacea clones was investigated. In this experiment clones were grown from two connected parent ramets and the parts of the clone produced by each parent ramet were subjected independently to either nutrient-rich or nutrient-poor conditions. Ramets in resource-rich conditions provided considerable physiological support to those in resource-poor conditions. This was measured as a dry weight gain compared with the weight of the corresponding part of the control clones growing in resource-poor conditions. However, when stolons grew from resource-poor conditions into resource-rich conditions, there was no similar evidence of the resourcepoor ramtes receiving support from resource-rich ramets. Physiological integration did not result in dry weight gains when this would have necessitated basipetal translocation of resources.Because of the predominantly acropedal direction of movement of translocates in G. hederacea, the structure of the clone was important in determining the effectiveness of integration between ramets. Where physiological integration was effective, the cost to the supporting ramets in terms of dry weight was insignificant. Physiological integration allows clones to maintain a presence in less favourable sites with insignificant cost to ramets in favourable sites, thereby reducing the probability of invasion by other plants, and providing the potential for rapid clonal growth if conditions improve. Integrated support of ramets in unfavourable conditions also enables the clone to grow through unfavourable sites, thus increasing the probability of encountering more favourable conditions by wider foraging.     

Low plant density enhances gene dispersal in the Amazonian understory herb Heliconia acuminata

In theory, conservation genetics predicts that forest fragmentation will reduce gene dispersal, but in practice, genetic and ecological processes are also dependent on other population characteristics. We used Bayesian genetic analyses to characterize parentage and propagule dispersal in Heliconia acuminata L. C. Richard (Heliconiaceae), a common Amazonian understory plant that is pollinated and dispersed by birds. We studied these processes in two continuous forest sites and three 1‐ha fragments in Brazil's Biological Dynamics of Forest Fragments Project. These sites showed variation in the density of H. acuminata. Ten microsatellite markers were used to genotype flowering adults and seedling recruits and to quantify realized pollen and seed dispersal distances, immigration of propagules from outside populations, and reproductive dominance among parents. We tested whether gene dispersal is more dependent on fragmentation or density of reproductive plants. Low plant densities were associated with elevated immigration rates and greater propagule dispersal distances. Reproductive dominance among inside‐plot parents was higher for low‐density than for high‐density populations. Elevated local flower and fruit availability is probably leading to spatially more proximal bird foraging and propagule dispersal in areas with high density of reproductive plants. Nevertheless, genetic diversity, inbreeding coefficients and fine‐scale spatial genetic structure were similar across populations, despite differences in gene dispersal. This result may indicate that the opposing processes of longer dispersal events in low‐density populations vs. higher diversity of contributing parents in high‐density populations balance the resulting genetic outcomes and prevent genetic erosion in small populations and fragments.     

A transcontinental comparison of the diversity and composition of tropical forest understory herb assemblages

Although tropical forests are renowned for their high plant diversity, to date there has been no global quantitative evaluation of the local species richness of terrestrial forest herbs in tropical forests. In this paper, richness and composition of terrestrial herb assemblages is compared in tropical forests of America, Africa and South East Asia. We established 86 non-continuous transects of 445 m each. Herb species richness was analysed and compared to six environmental parameters using minimal adequate regression models and simultaneous autoregressive models. At the global scale, we found a close relationship between herb species richness and temperature parameters, with no differences between continents. The subdivision into three main taxonomic groups (ferns, monocots, dicots) showed that each group has distinct relations to environmental factors and differences in richness between continents. Most of the 72 families found have pantropical distributions but 12, 11, and 16 families were significantly over-represented in America, Africa, and Asia, respectively. Although total species richness was closely related to climatic factors, ferns, monocots and dicots were represented by distinct sets of families with varying species richness on each continent. Which species are found at a given site may thus reflect group-specific evolutionary and historical factors.     

Distribution of genetic diversity among disjunct populations of the rare forest understory herb, Trillium reliquum

We assessed genetic diversity and its distribution in the rare southeastern US forest understory species, Trillium reliquum. In all, 21 loci were polymorphic (PS=95.5%) and the mean number of alleles per polymorphic locus was 3.05. However, genetic diversity was relatively low (Hes=0.120) considering the level of polymorphism observed for this outcrossing species. A relatively high portion of the genetic diversity (29.7%) was distributed among populations. There was no relationship between population size and genetic diversity, and we did not detect significant inbreeding. These results are best explained by the apparent self-incompatibility of this species, its longevity and clonal reproduction. To address questions regarding the history of T. reliquum's rarity, we compared results for T. reliquum with that of its more common and partially sympatric congener, T. cuneatum. Despite shared life history traits and history of land use, we observed significant genetic differences between the two species. Although T. cuneatum contains slightly lower polymorphism (Ps=85%), we detected significantly higher genetic diversity (Hes=0.217); most of its genetic diversity is contained within its populations (GST=0.092). Our results suggest that not only is there little gene flow among extant T. reliquum populations, but that rarity and population isolation in this species is of ancient origins, rather than due to more recent anthropogenic fragmentation following European colonization. The Chattahoochee River was identified as a major barrier to gene exchange.     

Effects of forest age and disturbance on population persistence in the understory herb,Arisaema triphyllum (Araceae)

Long-lived understory herbs experience a highly dynamic forest over space and time, yet can persist for more than a century. To understand how these populations persist, we examined effects of forest age and disturbance on potential sexual reproduction and clonal growth in the sexually labile perennial, Arisaema triphyllum. Potential sexual reproduction (female:male ratio) was significantly greater in the Young and Old-Gap forest states compared with Old, closed-canopy sites, where it was virtually absent. In contrast, clonal growth (estimated by cormlet production) did not differ significantly among the three forest states. Of seven environmental variables measured, only light (positively) and plant density (negatively) contributed significantly to the variation in potential sexual reproduction, while no measured variables contributed significantly to the variation in number of cormlets. The larger sexual reproductive effort (flower+stalk biomass/total biomass) for males in the undisturbed, 100 yr old forest may explain the absence of females in these sites, while the invariant vegetative reproductive effort (cormlet biomass/total biomass) may explain the similarity in average number of cormlets per individual per season across forest states. These results suggest that potential sexual reproduction is resource-limited, while clonal growth may be resource-independent. By maintaining ramet production during unfavorable periods, A. triphyllum populations disperse temporally, waiting for conditions under which sexual reproduction may resume.     

Heavy metal accumulation and distribution in forest understory herb species of Carpathian beech ecosystems

Abstract

This study addresses the issue of heavy metal (HM) accumulation and distribution for three different plant species, Carex pilosa, Dentaria bulbifera, Galium odoratum, in Carpathian beech ecosystems. Data are presented on HM concentrations in forest understory vegetation and a preliminary insight into different HM allocation patterns is provided. Bioaccumulation factors (BCFs) and shoot/root ratios differed considerably among the species and between polluted and unpolluted regions. HMs were accumulated in forest plants as follows: Cu > Zn > Cd >Pb in unpolluted areas and Zn> Cd > Cu >Pb in polluted areas. Zn was preferentially distributed to roots and Cu to shoots. The distribution of Cd and Pb in different plant parts was specific in terms of the species-dependence. Cd and Pb levels in Carex pilosaand Galium odoratumwere more strictly controlled in the transfer zone of root-shoot, compared to Dentaria bulbifera.The highest BCFs were found in Carex pilosafor Cu (5.9) and in Dentaria bulbiferawas found the highest shoot/root ratio for Cd (3.1).     

Sources of variation in the nesting success of understory tropical birds

Survival of offspring is a key fitness component and, for birds, the threat of predation on nests is especially influential. Data on rates of nest success from tropical regions are comparatively few, conservation‐relevant, and essential for assessing the validity of models comparing the life histories and behavior or birds across latitudinal gradients. We monitored over 2 000 nests in the lowland forests of central Panama and, using the logistic exposure to model the fate of nests, explored the importance of variation in rate of nest success according to type of nest, height of nests, among years, in early versus late nests, and at different stages of the nest cycle. Analyses of over 1 400 nests for 18 species revealed considerable variation among species in the daily survival rate of nests (range among 18 species=0.91 to 0.98), but nest type and stage of the nesting cycle were generally influential on the probability of nest success. Cavity or enclosed nesters experienced greater nest success than open cup nesters and rates of nest loss were generally greatest in the nestling stage. We found limited evidence that height of nests affected probability of success, but no indication that timing of nesting effort was influential. Despite the occurrence of a severe ENSO event during our sampling, annual variation in nest success was not consistent among species. Interspecific variation in the rates and patterns of nest predation in our study, coupled with reports of high rates of nest loss at temperate latitudes, lead us to question long standing assumptions about latitudinal trends in rates of nest loss. We urge further work to understand the implications of nest predation on the evolutionary ecology of tropical birds.     

The dynamics of antigenic variation and growth of African trypanosomes

Antigenic variation in African trypanosomes, which is a simple strategy for survival in the immune host, is rendered complex by its magnitude. For protection from nonspecific immunity and escape from specific immunity, each trypanosome is covered by a replaceable surface coat composed of the variant surface glycoprotein (VSG), which specifies the variable antigen type (VAT) of the trypanosome. Antigenic variation is the process by which the trypanosome switches from one coat to another. Here, David Barry and Michael Turner consider this phenomenon within the context of the course of trypanosome infection.