Variation in woody plant δ13C along a topoedaphic gradient in a subtropical savanna parkland

δ¹³C values of C₃ plants are indicators of plant carbon–water relations that integrate plant responses to environmental conditions. However, few studies have quantified spatial variation in plant δ¹³C at the landscape scale. We determined variation in leaf δ¹³C, leaf nitrogen per leaf area (N_area), and specific leaf area (SLA) in April and August 2005 for all individuals of three common woody species within a 308 × 12-m belt transect spanning an upland–lowland topoedaphic gradient in a subtropical savanna in southern Texas. Clay content, available soil moisture, and soil total N were all negatively correlated with elevation. The δ¹³C values of Prosopis glandulosa (deciduous N₂-fixing tree legume), Condalia hookeri (evergreen shrub), and Zanthoxylum fagara (evergreen shrub) leaves increased 1–4‰ with decreasing elevation, with the δ¹³C value of P. glandulosa leaves being 1–3‰ higher than those of the two shrub species. Contrary to theory and results from previous studies, δ¹³C values were highest where soil water was most available, suggesting that some other variable was overriding or interacting with water availability. Leaf N_area was positively correlated with leaf δ¹³C of all species (p < 0.01) and appeared to exert the strongest control over δ¹³C along this topoedaphic gradient. Since leaf N_area is positively related to photosynthetic capacity, plants with high leaf N_area are likely to have low p _I/p _a ratios and therefore higher δ¹³C values, assuming stomatal conductance is constant. Specific leaf area was not correlated significantly with leaf δ¹³C. Following a progressive growing season drought in July/August, leaf δ¹³C decreased. The lower δ¹³C in August may reflect the accumulation of ¹³C-depleted epicuticular leaf wax. We suggest control of leaf δ¹³C along this topoedaphic gradient is mediated by leaf N_area rather than by stomatal conductance limitations associated with water availability.     

Does competition or facilitation regulate migrant ungulate populations in the Serengeti? A test of hypotheses

Summary Interspecific competition and facilitation have both been proposed as processes promoting species separation and co-existence in African ungulates. In one group of grazers on the Serengeti plains, comprising wildebeest (Connochaetes taurinus), zebra (Equus burchelli), and Thomson's gazelle (Gazella thomsoni), these processes have also been suggested to regulate the populations. Censuses of these populations over 20 years have shown changes that allow a test of which, if either, process regulates population numbers. Wildebeest numbers have levelled off as a result of intraspecific competition for food following a five-fold increase due to release from disease and an increase in food supply. Zebra numbers have remained stationary throughout the same period. Gazelle numbers have declined in the last 10 years. These results are contrary to the facilitation hypothesis, which predicts that wildebeest numbers should not have increased if those of zebra did not, and that gazelle numbers should have increased since the wildebeest population increased. The gazelle results are consistent with the interspecific competition hypothesis, but the zebra results are contrary to it. We propose an alternative hypothesis that predation regulates the zebra population, and we suggest ways of testing this.     

Soil moisture redistribution as a mechanism of facilitation in savanna tree–shrub clusters

Plant–soil water relations were examined in the context of a selective removal study conducted in tree–shrub communities occupying different but contiguous soil types (small discrete clusters on shallow, duplex soils versus larger, extensive groves on deep, sandy soils) in a subtropical savanna parkland. We (1) tested for the occurrence of soil moisture redistribution by hydraulic lift (HL), (2) determined the influence of edaphic factors on HL, and (3) evaluated the significance of HL for overstory tree–understory shrub interactions. Diel cycling and nocturnal increases in soil water potential (Ψ_soil), characteristic signatures of HL, occurred intermittently throughout an annual growth cycle in both communities over a range of moisture levels (Ψ_soil=−0.5 to −6.0 MPa) but only when soils were distinctly stratified with depth (dry surface/wet deep soil layers). The magnitude of mean (±SE) diel fluctuations in Ψ_soil (0.19±0.01 MPa) did not differ on the two community types, though HL occurred more frequently in groves (deep soils) than clusters (shallow soils). Selective removal of either Prosopis glandulosa overstory or mixed-species shrub understory reduced the frequency of HL, indicating that Prosopis and at least one other woody species was conducting HL. For Zanthoxylum fagara, a shallow-rooted understory shrub, Prosopis removal from clusters decreased leaf water potential (Ψ_leaf) and net CO₂ exchange (A) during periods of HL. In contrast, overstory removal had neutral to positive effects on more deeply-rooted shrub species (Berberis trifoliolata and Condalia hookeri). Removal of the shrub understory in groves increased A in the overstory Prosopis. Results indicate the following: (a) HL is common but temporally dynamic in these savanna tree–shrub communities; (b) edaphic factors influencing the degree of overstory/understory development, rooting patterns and soil moisture distribution influence HL; (c) net interactions between overstory and understory elements in these woody patches can be positive, negative and neutral over an annual cycle, and (d) Prosopis-mediated HL is an important mechanism of faciliation for some, but not all, understory shrubs.     

Effect of zooid spacing on bryozoan feeding success: is competition or facilitation more important?

Most Recent bryozoan species are encrusting sheets, and many of these colonies have densely packed feeding zooids. In this study, I tested whether tight packing of feeding zooids affects food capture. Colonies of a bryozoan with an encrusting sheet form (Membranipora membranacea) were dissected to produce individuals whose feeding zooids were (1) closely packed, (2) more widely spaced, or (3) isolated. For each type, rates of particle ingestion were measured in still water and in a freestream velocity of 2.7 cm s(-1). Ingestion rate increased when zooids were closest together, probably because of reduced refiltration and increased feeding current strength farther from the lophophores. The mean incurrent velocity within 0.02 cm above the center of the lophophore was 0.28 cm s(-1) regardless of zooid spacing; however, when the incurrent velocity was measured more than 0.1 cm from the lophophores, zooids that were close together or spaced one zooid's width apart had significantly faster incurrent velocities than single zooids. Flow visualization suggests that isolated zooids and those spaced far apart refilter more water than zooids that are close together. These results along with the observed trend of increased zooid integration over evolutionary time suggest that the benefits of increasing coordination outweigh the consequences of intrazooid competition.     

Centrosome competition: a possibility?

A recent theory on the evolution of sexuality, has hypothesized heritable variation in the functional properties of centrosomes, leading to competition for the organization of the mitotic spindle when different centrosomes enter a common cytoplasm. We present here data on polyethylene glycol-induced polykaryocytes of cultured Chinese hamster fibroblasts indirectly supporting centrosome competition. On the assumption that sib centrosomes are similar and variation increases with cell generations, the frequencies of multipolar mitoses were compared in cultures fused under conditions favoring sib cell fusion or fusion of distantly related cells. Multipolar mitoses were considerably more frequent in the former, when the average difference between pairs of centrosomes was assumed to be too small for one centrosome to "outcompete" the other.     

ABA and sugar interactions regulating development: cross-talk or voices in a crowd?

Plant growth and development are controlled by the concerted action of many signaling pathways that integrate information from environmental signals with that from developmental and metabolic cues. Physiological studies have demonstrated that abscisic acid and sugars have both similar and antagonistic effects on diverse processes, including seed development, germination, and seedling growth. Recent genetic studies have identified several loci that are involved in both sugar and hormonal responses. It is rarely clear whether these apparent linkages reflect direct or indirect interactions between sugar and hormone signaling pathways, but the identification of gene products that are encoded at these loci is allowing these possibilities to be tested.     

What limits herb biomass in grasslands: competition or herbivory?

Competition and herbivory are two of the main forces shaping plant communities. Although several studies have investigated their impact on plant populations separately, few investigations have examined how they might interact. With the purpose of clarifying the combined roles of competition and herbivory on herb biomass in a grassland, we assessed the effects of different herbivores with reduced grass competition. We conducted a field experiment in 2000-2001 in a British acid grassland (Oak Mead), where we experimentally reduced grass biomass and excluded rabbits, insects and molluscs in a factorial design. Removing the grasses from Oak Mead dramatically increased herb biomass and total above-ground biomass. Herbivore exclusions (i.e. rabbits, insects and molluscs) did not affect total above-ground biomass, but they altered relative abundance of several species. Grass removal and rabbit exclusion had positive interactions on biomass of several herb species, and there were some subtle interactions between different herbivore groups: monocots benefitted when both rabbits and molluscs were excluded, and some herb species had greater biomass when insects and rabbits were absent. We then compared the results with a 10-year experiment that manipulated similar variables in neighbouring grassland (Nashs Field). The comparison between Oak Mead and Nashs Field showed that cessation of herbicide application returns the system to its previous state of grass dominance after 3 years. Therefore, even when herbs were more abundant, they could not prevent reinvasion of the grasses once external factors were removed.     

Evidence of a tipping point in a southern African savanna?

Many ecosystems exhibit threshold behaviour, where periods of relative stability are punctuated by rapid transitions between alternate stable states when an ecological threshold, or tipping point, is reached. This is of concern in grass-dominated habitats, many of which appear to be on the point of conversion to more wooded vegetation assemblages. However, changes in grass-dominated ecosystems are often difficult to interpret, because it is not always clear whether grasslands are ancient or are anthropogenically derived from past deforestation. As a result, the conservation, maintenance and restoration of ancient grasslands are sometimes neglected.In this study, the history of vegetation change in the savannas of the Hluhluwe-iMfolozi Park, KwaZulu-Natal, South Africa, are investigated by analysing stable carbon isotopes (δ¹³C) from soil profiles. Without exception, the data show that C₃ dominated thicket, forest, and densely wooded savanna now occur on sites that were previously C₄ grassland or open savanna. Although the drivers of this change are not clear, there is potential for management intervention because tree density can be manipulated through fire, a natural part of this dynamic landscape. The study identified two sites which are at a threshold between C₄ and C₃ dominance, and highlighted them as priorities for conservation management intervention.     

Gene-environment interactions in human disease: nuisance or opportunity?

Many environmental risk factors for common, complex human diseases have been revealed by epidemiologic studies, but how genotypes at specific loci modulate individual responses to environmental risk factors is largely unknown. Gene-environment interactions will be missed in genome-wide association studies and could account for some of the 'missing heritability' for these diseases. In this review, we focus on asthma as a model disease for studying gene-environment interactions because of relatively large numbers of candidate gene-environment interactions with asthma risk in the literature. Identifying these interactions using genome-wide approaches poses formidable methodological problems, and elucidating molecular mechanisms for these interactions has been challenging. We suggest that studying gene-environment interactions in animal models, although more tractable, might not be sufficient to shed light on the genetic architecture of human diseases. Lastly, we propose avenues for future studies to find gene-environment interactions.     

Is the balance between competition and facilitation a driver of the patch dynamics in arid vegetation mosaics?

In most arid ecosystems, the vegetation is organized into two‐phase mosaics, where high‐cover vegetation patches are interspersed in a matrix of low plant cover. We studied the role of the biotic interaction balance (competition/facilitation) between shrubs and grasses as a driver of patch dynamics and maintenance of two‐phase vegetation mosaics. Following Watt's seminal model, we conducted two field experiments in which we manipulated different vegetation patches to obtain the different stages along the building and degradation dynamics of high‐cover patches. In addition we applied two possible belowground competition treatments (natural and experimentally reduced). We measured plant variables (emergence, survival, height, flower culms) on grass seedlings and transplants. We integrated all plant measurements into a single positive and negative component, to calculate the net balance along three stages of the patch dynamics proposed. The net biotic interaction balance was negative during the early and mature stages of high‐cover patches because the average standardized effect from the negative component was below ?0.44, while the positive component was not different from zero. However, the net biotic interaction balance was positive during the degraded stages of high‐cover patches because the negative average component was ?0.37, while the positive component reached 0.58. The negative net effects during early and mature stages of high‐cover patches can be explained by the occurrence of wet years, because high rainfalls hide the aboveground facilitation. Our findings point out the importance of complementary mechanisms to the interaction balance in the mosaic maintenance (e.g. trapping of seeds by shrubs) according to the inter‐annual rainfall variability.     

Rainforest and savanna landscape dynamics in New Caledonia: Towards a mosaic of stable rainforest and savanna states?

Stable forested environments can be converted to savanna in response to changes in environmental disturbances. New Caledonia is a biodiversity hotspot; significant ecological and economic resources would be lost if forests were turned into savanna by anthropogenic environmental changes. On the landscape scale, systems that have undergone shifts of this kind are characterized by sharp forest–savanna boundaries and mosaic‐like distributions of savanna and forest. Understanding the locations and the dynamics of such boundaries is a challenge for ecologists and is critical for landscape management and biodiversity conservation. Using a time series of aerial photographs (1955–2000) and a forest habitat suitability map, we tested the hypothesis that topography and spatial processes, especially those relating to fire spread and seed dispersal, are the main determinants of the spatial distribution of rainforest and savanna in a New Caledonian landscape covering 24 km². Within the studied landscape, the overall forest coverage decreased by 24% between 1976 and 2000. This was primarily due to the contraction of forests on west‐facing slopes, which accounted for about 90% of the total loss. Conversely, the east‐facing forests seemed to have contracted extensively prior to the studied period, and were confined to refuges. A habitat suitability index calculated from the landscape's topographical features using generalized additive models accurately predicted both the presence of forests and the probability of forest expansion/contraction. We also provide evidence that spatial processes such as fire spread and seed dispersal limit the expansion and contraction of forests. Our results suggest that rainforests on west‐facing slopes in New Caledonia will be progressively destroyed by fire until they are restricted to refuges along thalwegs and creeks, as appears to have already happened for their east‐facing counterparts.     

Water use by woody plants on contrasting soils in a savanna parkland: assessment with δ2H and δ18O

In savanna parklands of southern Texas, patches of grassland and discrete clusters of small trees and shrubs occur on sandy loam surface soils underlain by an argillic horizon (claypan) at 40 cm. Large trees and shrubs in groves occur on deep (2 m) sandy loam soils without an argillic horizon. 2H and 18O of rainfall, groundwater, and soil and plant water were measured to: (1) determine if coexistence in woody patches occurs via vertical stratification of soil water uptake; (2) document differences in plant water acquisition on contrasting soil types; and (3) evaluate recharge and evaporative losses of soil moisture from grassland vs. wooded landscape elements. Groundwater was isotopically similar to weighted rainfall, suggesting local recharge at this site. Linear regressions of soil water 2H on 18O yielded slopes less than the meteoric water line, indicating significant evaporative losses of soil moisture in all landscape elements. Interspecific differences in root density distribution were significant; some woody species had roots well below 1.6 m, while others had few roots below 0.8 m. 2H and 18O values of stem water from all plants in groves were lower than those of soil water in the upper 1.5 m of the profile, suggesting all species obtained their water from depths >1.5 m. Deep roots of trees and shrubs at this savanna parkland site thus appeared to have a functional significance that was not revealed by biomass or density determinations. Root densities of species in discrete clusters (claypan present) were typically greater than those of the same species in groves (claypan absent), especially in the upper 80 cm of the soil profile. Consistent with rooting profiles, 2H and 18O values of plant water indicated that trees and shrubs in discrete clusters with fine- textured subsoils obtained most of their water at depths <1.5 m. As with groves, there was no indication of water resource partitioning between species. In summary, we saw no isotopic evidence that co- occurring woody plants at this savanna parkland site were partitioning soil moisture vertically during late summer/early fall, despite marked differences in their root density distributions. This supports other lines of evidence which indicate that species interactions in tree/shrub clumps are competitive, and that species composition is therefore unstable in those landscape elements.     

Porphyrin-membrane interactions: binding or partition?

Porphyrins are photodynamic drugs employed in an experimental tumor-treatment modality in which cell membranes are one of the primary drug-action sites. To gain insight into the nature of the interaction of these drugs with those primary sites we have studied the affinity of porphyrins to the lipid moieties of biological membranes, at the molecular level. The association of porphyrins to large unilamellar liposomes, modeling the lipid regions of biological membranes was studied (at equilibrium) for deuteroporphyrin IX and protoporphyrin IX, at neutral pH and 37 degrees C, taking into account porphyrin aggregation. Two thermodynamic approaches were investigated: (i) Simple partition equilibria between the external aqueous phase and the lipid bilayer, for drug monomers and dimers. (ii) Binding equilibria of drug monomers and dimers to the lipid bilayer. Using two types of experimental design and processing the data according to the expectations of both approaches, three different models for the binding (differing in the participation assigned to the dimer) were considered. Our major findings are: (a) The data clearly do not fit with the expectations for simple partition equilibria, nor with binding models assuming direct participation of the dimers. (b) The data fit well with a binding process, in which the membrane binds the porphyrin monomers only, with the dimers participating indirectly through the aqueous dimerization equilibrium. (c) At 37 degrees C and neutral pH, for liposomes composed of phosphatidylcholine/cholesterol at molar ratios of 3:2, we found for both investigated species a binding constant of 2.3 x 10(4) M-1. (d) For each species the binding constant is independent of the initial and final states of drug aggregation in the aqueous phase.     

Nomenclature: or what's in a name?

Transgenic Research -     

Wind or insect pollination? Ambophily in a subtropical gymnosperm Gnetum parvifolium (Gnetales)

Gnetum (Gnetales) has long been regarded as insect pollinated due to its range‐restricted distribution in tropical rain forests, where wind pollination is supposed to be detrimental. However, ubiquitous pollen limitation in the tropics might cause transition to anemophily, or even ambophily for reproductive assurance, especially for gymnosperms such as Gnetum, which exhibit various anemophilous syndromes, including absence of petals, greenish color of strobili, dioecy and uniovulate flowers. Pollination treatments applied to Gnetum parvifolium in southern China revealed this rare and endangered species to be pollen limited and incapable of apomixis. Pollen grains of G. parvifolium did not adhere to each other and could be trapped by interception traps near the male and female strobili. Seed set in the netted treatment (anemophily) was significantly higher than in the bagged treatment (apomixis) but lower than open pollination (anemophily and entomophily), indicating that both wind and biotic pollination contributed to seed production of G. parvifolium. The occurrence of ambophily in Gnetum and the prevalence of anemophily in Ehpedra and Welwitschia suggest that wind may also play a role in the pollination of the ancient Gnetales.     

Intraspecific competition replaces interspecific facilitation as abiotic stress decreases: The shifting nature of plant–plant interactions

Plant–plant interactions change depending on environmental conditions, shifting from competition to facilitation when the stress is high. In addition to these changes, the relevance of intraspecific compared to interspecific interactions may also shift as abiotic stress does. We inferred intra- and interspecific plant–plant interactions of the cushion plant Hormathophylla spinosa as related to the dominant shrub Juniperus sabina in two sites with contrasting abiotic conditions (a slope with high-stress conditions vs. a valley bottom with milder conditions) in a Mediterranean high mountain. Specifically, we studied the spatial patterns and several variables related to plant performance (plant size and form, non-structural carbohydrate – NSC – concentrations and radial growth) of H. spinosa.The spatial pattern varied depending on site conditions. H. spinosa plants were positively associated with juniper in the high-stress slope site, probably through higher establishment rates due to the amelioration of soil conditions. In contrast, in the milder valley site H. spinosa establishment occurred mostly in open areas. Age structure, inferred from annual rings, reflected a massive establishment event in the whole study area which occurred 30–50 years ago. Canopy variables and radial growth were density dependent: both were negatively affected by the high density of H. spinosa individuals in the valley, but favoured by junipers on the slope. Interestingly, NSCs showed the opposite pattern, suggesting lower investment in growth by H. spinosa plants in the valley than on the slope.Our results reinforce the strong links existing between intra- and interspecific relationships and the need to include both when studying the influence of abiotic conditions on plant–plant interactions. This approach enabled us to detect that the direction and intensity of plant–plant interactions may shift at different ecological levels. Particularly interesting was the finding that optimal sites at the population level may not necessarily be the sites showing maximum individual performance.     

Submerged macrophyte zonation in a Mediterranean salt marsh: a facilitation effect from established helophytes?

Abstract. The relationship between communities of submerged annual macrophytes (predominately Chara spp.) and environmental characteristics is studied in three habitats with different dominant perennial species (Arthrocnemum, Juncus, Scirpus) and areas of bare soil. The distribution of submerged annual macrophytes is significantly dependent on two independent environmental factors: a dominant gradient of flooding/salinity, and a secondary gradient of nutrients related exclusively to the combined abundance of helophytes of the genus Scirpus (S. maritimus and S. litoralis). The results suggest that these emergent helophytes (1) are able to modify water column parameters (such as concentration of nitrates, phosphates, potassium, and bicarbonates) which are important for the communities of submerged macrophytes, and (2) play a fundamental role in the generation of secondary sources of environmental variability which, superimposed on the main gradient of flooding/salinity, favours the appearance of new compositional equilibria in such communities. The existence of a process of facilitation is discussed by which the emergent helophytes induce changes in nutrient availability that would favour relatively nutrient‐demanding charophyte species (such as Chara connivens and Nitella hyalina), altering the established relationships with other coexisting charophytes (such as Chara canescens and C. galioides) that dominate in the absence of the facilitating species. Nevertheless, the increased nutrient concentration associated to the presence of helophytes would not introduce significant changes in the total biomass of submerged aquatic macrophytes.