Effects of plant nutrition on the balance of insect relevant cardenolides and glucosinolates in Erysimum cheiranthoides

The possible effects of environmental stress on plant chemistry that are important to herbivorous insects were examined by growing a wild crucifer, Erysimum cheiranthoides, under different nutrient regimes. Oviposition by the cabbage butterfly, Pieris rapae, is thought to be affected by the balance of glucosinolates (stimulants) and cardenolides (deterrents) at the surface of leaves. E. cheiranthoides seedlings were provided with three levels of nitrogen and two levels of sulfur for a period of 15 days before analysis of semiochemicals in whole leaf tissue and at the surface of the foliage. The ratio of cardenolides to glucosinolates in the plants at elevated C/N ratios followed the carbon/nutrient balance hypothesis. However, a high nitrogen supply enhanced biomass production to the extent that concentrations of secondary compounds were unchanged or reduced. The concentration of glucosinolates (glucoiberin and glucocheirolin) at the surface was positively related to whole tissue levels. However, cardenolide (erysimoside and erychroside) concentrations, which were highest in leaf tissue of nitrogen-deficient plants, had the lowest surface levels on foliage of these plants. Possible reasons for differential expression of cardenolides and glucosinolates in a plant as a result of nutrient deficiency are discussed.     

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.     

Belowground positive and negative feedbacks on CO2 growth enhancement

In this paper we present a conceptual model of integrated plant-soil interactions which illustrates the importance of identifying the primary belowground feedbacks, both positive and negative, which can simultaneously affect plant growth responses to elevated CO₂. The primary negative feedbacks share the common feature of reducing the amount of nutrients available to plants. These negative feedbacks include increased litter C/N ratios, and therefore reduced mineralization rates, increased immobilization of available nutrients by a larger soil microbial pool, and increased storage of nutrients in plant biomass and detritus due to increases in net primary productivity (NPP). Most of the primary positive feedbacks share the common feature of being plant mediated feedbacks, the only exception being Zak et al.'s hypothesis that increased microbial biomass will be accompanied by increased mineralization rates. Plant nutrient uptake may be increased through alterations in root architecture, physiology, or mycorrhizal symbioses. Further, the increased C/N ratios of plant tissue mean that a given level of NPP can be achieved with a smaller supply of nitrogen.Identification of the net plant-soil feedbacks to enhanced productivity with elevated CO₂ are a critical first step for any ecosystem. It is necessary, however, that we first identify how universally applicable the results are from one study of one ecosystem before ecosystem models incorporate this information. The effect of elevated CO₂ on plant growth (including NPP, tissue quality, root architecture, mycorrhizal symbioses) can vary greatly for different species and environmental conditions. Therefore it is reasonable to expect that different ecosystems will show different patterns of interacting positive and negative feedbacks within the plant-soil system. This inter-ecosystem variability in the potential for long-term growth responses to rising CO₂ levels implies that we need to parameterize mechanistic models of the impact of elevated CO₂ on ecosystem productivity using a detailed understanding of each ecosystem of interest.