Topcats and underdogs: intraguild interactions among three apex carnivores across Asia's forestscapes

Intraguild interactions among carnivores have long held the fascination of ecologists. Ranging from competition to facilitation and coexistence, these interactions and their complex interplay influence everything from species persistence to ecosystem functioning. Yet, the patterns and pathways of such interactions are far from understood in tropical forest systems, particularly across countries in the Global South. Here, we examined the determinants and consequences of competitive interactions between dholes Cuon alpinus and the two large felids (leopards Panthera pardus and tigers Panthera tigris) with which they most commonly co-occur across Asia. Using a combination of traditional and novel data sources (N = 118), we integrate information from spatial, temporal, and dietary niche dimensions. These three species have faced catastrophic declines in their extent of co-occurrence over the past century; most of their source populations are now confined to Protected Areas. Analysis of dyadic interactions between species pairs showed a clear social hierarchy. Tigers were dominant over dholes, although pack strength in dholes helped ameliorate some of these effects; leopards were subordinate to dholes. Population-level spatio-temporal interactions assessed at 25 locations across Asia did not show a clear pattern of overlap or avoidance between species pairs. Diet-profile assessments indicated that wild ungulate biomass consumption by tigers was highest, while leopards consumed more primate and livestock prey as compared to their co-predators. In terms of prey offtake (ratio of wild prey biomass consumed to biomass available), the three species together harvested 0.4–30.2% of available prey, with the highest offtake recorded from the location where the carnivores reach very high densities. When re-examined in the context of prey availability and offtake, locations with low wild prey availability showed spatial avoidance and temporal overlap among the carnivore pairs, and locations with high wild prey availability showed spatial overlap and temporal segregation. Based on these observations, we make predictions for 40 Protected Areas in India where temporally synchronous estimates of predator and prey densities are available. We expect that low prey availability will lead to higher competition, and in extreme cases, to the complete exclusion of one or more species. In Protected Areas with high prey availability, we expect intraguild coexistence and conspecific competition among carnivores, with spill-over to forest-edge habitats and subsequent prey-switching to livestock. We stress that dhole–leopard–tiger co-occurrence across their range is facilitated through an intricate yet fragile balance between prey availability, and intraguild and conspecific competition. Data gaps and limitations notwithstanding, our study shows how insights from fundamental ecology can be of immense utility for applied aspects like large predator conservation and management of human–carnivore interactions. Our findings also highlight potential avenues for future research on tropical carnivores that can broaden current understanding of intraguild competition in forest systems of Asia and beyond.     

Characteristics and Regulatory Properties of the H+-ATPase in a Plasma Membrane Fraction Purified from Arabidopsis thaliana

The aqueous two-phase partitioning technique was utilized to isolate a plasma membrane (PM) fraction from etiolated seedlings of Arabidopsis thaliana. The purification procedure adopted yielded a fraction highly enriched in PM as compared to inner membranes, with a recovery of about 30%, as judged from the activities of PM markers such as vanadate-sensitive ATPase, FC binding and UDP-glucose sterol glucosyltransferase. The purified PM fraction displayed vanadate-sensitive H⁺ pumping activity. Its purity was confirmed by the biochemical characteristics of its ATPase activity assayed in the absence of Ca²⁺: sensitivity to vanadate (IC₅₀ ca. 1 μM), Mg²⁺-dependence, insensitivity to molybdate, oligomycin and nitrate, pH optimum at 6.6. The PM H⁺-ATPase activity was stimulated by fusicoccin and by a controlled treatment of the PM with trypsin. In both cases stimulation was much stronger on the activity assayed at pH 7.5 than on the activity at pH 6.6. Moreover, neither fusicoccin nor the treatment with trypsin stimulated the portion of activity (30 to 40% at pH 7.5) which decayed upon preincubation of the PM in assay medium without ATP.     

PHOTOADAPTATION AND THE “PACKAGE” EFFECT IN DUNALIELLA TERTIOLECTA (CHLOROPHYCEAE)1

In the marine unicellular chlorophyte, Dunaliella tertiolecta Butcher, the spectrally averaged m vivo absorption cross section, normalized to chlorophyll a (so-called a* values), vary two-fold in response to changes in growth irradiance. We used a kinetic approach to examine the specific factors which account for these changes in optical properties as cells photoadapt. Using Triton X-100 to solubilize membranes, we were able to differentiate between “package” effects and pigmentation effects. Our analyses suggest that 43–49% of the variability in a* is due to changes in pigmentation, whereas 51–57% is due to the “package” effect. Further analyses revealed that changes in cell sue did not significantly affect packaging, while thylakoid stacking and the transparency of thylakoid membranes were important factors. Our results suggest that thylakoid membrane protein/lipid ratios change during photoadaptation, and these changes influence the effective rate of light harvesting per unit chlorophyll a.     

Sectoriality and physiological organisation in herbaceous plants: an overview

The physiological organisation of plants is considered in relation to the carbon economy of plant parts. Although assimilate is partitioned according to the relative strength of sinks, in many species there is also a very close relationship between partitioning and shoot phyllotaxy, giving rise to sectorial patterns of allocation whereby only certain sinks are supported by any source leaf. Essentially these sinks are in the same orthostichy as the source leaf. This constraint of the vascular architecture on assimilate distribution to developing sinks such as leaves, flowers and fruits is not always absolute, as following the loss of their principal source leaves these sinks can in many cases be supplied with assimilate by other leaves via new inter-orthostichy pathways. The supply of assimilate to major sinks such as developing fruits becomes more and more localised with time so that a fruit in an axillary position becomes largely supported by its subtending leaf; the reproductive node—a metamer-can thus be regarded as a relatively autonomous unit of the plant (an IPU). Similary, once established after a developmental phase of assimilate import, tiller ramets and branches in unitary plants tend to become physiologically autonomous modules. However, the functional autonomy of tillers is reversed following defoliation or shading as they are then sustained by the import of assimilate, subject to its availability, from unaffected tillers. Consequently the plant becomes physiologically integrated by the flow of assimilate from one part to another. The mainly autonomous ramets of many stoloniferous and rhizomatous species display a similar pattern of physiological integration in response to source manipulation, but in some species the ramets appear to maintain their independent functioning as a normal feature of the carbon allocation within the clone. In other clonal species, as the clone develops and becomes more structurally complex, vascular constraints start to restrict the movement of resources, and the clone becomes composed of a number of semi-autonomous IPUs. In unitary plants branches appear to remain very physiologically isolated in terms of their carbon economy once they become established, irrespective of a range of source-sink manipulations.These different patterns of physiological integration and organisation are discussed in relation to different strategies of assimilate utilisation and conservation.     

HMG-GoA reductase and terpenoid phytoalexins: Molecular specialization within a complex pathway

Terpenoid phytoalexins and other defense compounds play an important role in disease resistance in a variety of plant families but have been most widely studied in solanaceous species. The rate-limiting step in terpenoid phytoalexin production is mediated by 3-hydroxy-3-methylglutaryl coenzyme A reductase (HMGR), which catalyzes mevalonic acid synthesis. HMGRs are involved in the biosynthesis of a broad array of terpenoid compounds, and distinct isoforms of HMGR may be critical in directing the flux of pathway intermediates into specific end products. Plant HMGRs are encoded by a small gene family, and genomic or cDNA sequences encoding HMGR have been isolated from several plant species. In tomato, four genes encode HMGR; these genes are differentially activated during development and stress responses. One gene, hmg 2 , is activated in response to wounding and a variety of pathogenic agents suggesting a role in sesquiterpene phytoalexin biosynthesis. In contrast, expression patterns of tomato hmg l suggest a role in sterol biosynthesis and cell growth. Other plant species show an analogous separation of specific HMGR isoforms involved in growth and/or housekeeping function and inducible isoforms associated with biosynthesis of phytoalexins or other specialized "natural products". We are applying a variety of cell and molecular techniques to address whether subcellular localization and/or differential expression of these isoforms are key factors in determining end product accumulation during development and defense.     

Partitioning of water resources among plants of a lowland tropical forest

Source water used by plants of several species in a semi-evergreen lowland tropical forest on Barro Colorado Island, Panama, was assessed by comparing the relative abundance of deuterium, D, versus hydrogen, H (stable hydrogen isotope composition, D) in xylem sap and in soil water at different depths, during the dry season of 1992. Ecological correlates of source water were examined by comparing xylem water D values with leaf phenology, leaf water status determined with a pressure chamber, and rates of water use determined as mass flow of sap using the stem heat balance method. Soil water D values decreased sharply to 30 cm, then remained relatively constant with increasing depth. Average D values were-13, for 0–30 cm depth and-36.7 for 30–100 cm depth. Soil water D values were negatively associated with soil water content and soil water potential. Concurrent analyses of xylem water revealed a high degree of partitioning of water resources among species of this tropical forest. Xylem water D of deciduous trees (average=-25.3±1.4) was higher than that of evergreen trees (average=-36.3±3.5), indicating that evergreen species had access to the more abundant soil water at greater depth than deciduous species. In evergreen shade-tolerant and high-light requiring shrubs and small trees, D of xylem water was negatively correlated with transpiration rate and leaf water potential indicating that species using deeper, more abundant water resources had both higher rates of water use and more favorable leaf water status.     

Solubilization and hydrophobicity test by Triton X-114-partitioning of NADPH-protochlorophyllide oxidoreductase from the unicellular alga Scenedesmus obliquus, mutant C-2A'

NADPH-protochlorophyllide oxidoreductase (PChilde reductase, EC 1.3.1.33), a key enzyme in light-dependent greening and the conversion of etioplasts into chloroplasts was investigated in the the greening mutant C-2A' of the unicellular green alga Scenedesmus obliquus. In the absence of detergent, the solubilization of the enzyme increased with high glycerol concentrations in the buffer. Solubilization capacities of 4 non-ionic or zwitterionic detergents, Triton X-100, CHAPS, octylglucoside and decyl-maltopyranoside, were compared. Due to the addition of these detergents, the enzyme activity in the soluble fraction was increased severalfold. Hydrophobicity of the enzyme was analyzed by Triton X-114 phase partitioning. The protein had a preference for the aqueous phase, but its distribution was strongly influenced by the glycerol concentration of the buffer. These results indicate that the PChlide reductase of the green alga Scenedesmus obliquus is a hydrophobic, membrane-associated enzyme, but not an integral membrane protein.     

Root to shoot ratio of crops as influenced by CO2

Crops of tomorrow are likely to grow under higher levels of atmospheric CO₂. Fundamental crop growth processes will be affected and chief among these is carbon allocation. The root to shoot ratio (R:S, defined as dry weight of root biomass divided by dry weight of shoot biomass) depends upon the partitioning of photosynthate which may be influenced by environmental stimuli. Exposure of plant canopies to high CO₂ concentration often stimulates the growth of both shoot and root, but the question remains whether elevated atmospheric CO₂ concentration will affect roots and shoots of crop plants proportionally. Since elevated CO₂ can induce changes in plant structure and function, there may be differences in allocation between root and shoot, at least under some conditions. The effect of elevated atmospheric CO₂ on carbon allocation has yet to be fully elucidated, especially in the context of changing resource availability. Herein we review root to shoot allocation as affected by increased concentrations of atmospheric CO₂ and provide recommendations for further research. Review of the available literature shows substantial variation in R:S response for crop plants. In many cases (59.5%) R:S increased, in a very few (3.0%) remained unchanged, and in others (37.5%) decreased. The explanation for these differences probably resides in crop type, resource supply, and other experimental factors. Efforts to understand allocation under CO₂ enrichment will add substantially to the global change response data base.Abbreviations R:S root to shoot ratio, dry weight basis     

Geochemical partitioning and bioavailability of copper to aquatic plants in an artificial oxide-organic sediment

This study investigated the effects of competition between binding substrates (organic matter and iron oxide) and between metals (cadmium and copper), on the partitioning of sedimentary copper and its subsequent bioavailability to an aquatic plant. Organic matter and a synthesized iron oxide, ferrihydrite, were added singly and in combination to a series of sand sediments, which were then dosed with environmentally realistic concentrations of cadmium and copper and planted with rice,Oryza sativa. Organic matter controlled copper partitioning and bioavailability, whereas the synthetic ferrihydrite bound negligible amounts of either metal, even in the absence of organic matter. As organic matter concentrations increased, operationally-defined leachable copper decreased, organic-associated copper increased and the survival of rice plants improved in an approximately linear fashion. At a nominal starting copper concentration of 5.8 μg g dry wt⁻¹, plant survival after four weeks averaged 0–8% in sediments without organic matter, 25% in a sediment containing 0.18% organic matter and 58% in a sediment containing 0.36% organic matter. These results suggest that organic-associated forms of copper are unavailable to plants, and that the operational definition of ‘leachable’ copper (extracted with dilute ammonium acetate) adequately represents the species of copper that is (are) available to plants. Our study using a well-characterized artificial sediment supports the copper fractionation patterns and correlations between copper partitioning and bioavailability reported from the heterogeneous, poorly characterized sediments of natural lake and river sediments.     

Ontogenetic diet shifts and diet overlap among three closely related neotropical armoured catfishes

Analysis of stomach contents of larvae, juveniles and adults of three closely related armoured catfishes in coastal plain swamps of Suriname, South America, revealed major ontogenetic diet shifts. Larvae of Hoplosternum littorale, H. thoracatum and Callichthys callichthys showed a restricted diet of rotifers and microcrustacea. Diets of juveniles and adults were more diverse, but large volumes of microcrustacea, aquatic insect larvae and detritus were always present. In all three species diet overlap was especially low between larvae and adults. The proportion of detritus in the stomach was positively related to fish size, which probably reflected the inability of larger size classes to separate effectively benthic invertebrate prey from bottom substrata, and not a specialization towards detritivory. Adults showed only minor seasonality in diet composition, and no effect of season on diet breadth. Interspecific diet overlap among larvae, juveniles and adults did not differ significantly from 1 (complete overlap). Lack of food partitioning was probably related to a similar morphology of the three catfishes. Intraspecific between-size class diet overlap was considerably lower than interspecific within-size class overlap. It is suggested that large interspecific diet overlap occurred because food was not limiting the populations of these catfish. However, the nest habitats of the three species differed.