Differential utilization of summer rains by desert plants

Summary Seasonal changes in the hydrogen isotope ratios of xylem waters were measured to determine water sources used for growth in desert plants of southern Utah. While all species used winter-spring recharge precipitation for spring growth, utilization of summer rains was life-form dependent. Annuals and succulent perennials exhibited a complete dependence on summer precipitation. Herbaceous and woody perennial species simultaneously utilized both summer precipitation and remaining winter-spring precipitation, with herbaceous species much more reliant on the summer precipitation component. Several of the woody perennials exhibited no response to summer precipitation. Currently, precipitation in southern Utah is evenly partitioned between winter and summer time periods; however, global circulation models predict that summer precipitation will increase in response to anticipated climate change. Our data indicate that components within the community will differentially responde to the change in precipitation patterns. These results are discussed in relation to impact on competition and possible changes in community structure.     

Seasonal water uptake and movement in root systems of Australian phraeatophytic plants of dimorphic root morphology: a stable isotope investigation

A natural abundance hydrogen stable isotope technique was used to study seasonal changes in source water utilization and water movement in the xylem of dimorphic root systems and stem bases of several woody shrubs or trees in mediterranean-type ecosystems of south Western Australia. Samples collected from the native treeBanksia prionotes over 18 months indicated that shallow lateral roots and deeply penetrating tap (sinker) roots obtained water of different origins over the course of a winter-wet/summer-dry annual cycle. During the wet season lateral roots acquired water mostly by uptake of recent precipitation (rain water) contained within the upper soil layers, and tap roots derived water from the underlying water table. The shoot obtained a mixture of these two water sources. As the dry season approached dependence on recent rain water decreased while that on ground water increased. In high summer, shallow lateral roots remained well-hydrated and shoots well supplied with ground water taken up by the tap root. This enabled plants to continue transpiration and carbon assimilation and thus complete their seasonal extension growth during the long (4–6 month) dry season. Parallel studies of other native species and two plantation-grown species ofEucalyptus all demonstrated behavior similar to that ofB. prionotes. ForB. prionotes, there was a strong negative correlation between the percentage of water in the stem base of a plant which was derived from the tap root (ground water) and the amount of precipitation which fell at the site. These data suggested that during the dry season plants derive the majority of the water they use from deeper sources while in the wet season most of the water they use is derived from shallower sources supplied by lateral roots in the upper soil layers. The data collected in this study supported the notion that the dimorphic rooting habit can be advantageous for large woody species of floristically-rich, open, woodlands and heathlands where the acquisition of seasonally limited water is at a premium.     

Hydrogen and oxygen isotope ratios of tree-ring cellulose for riparian trees grown long-term under hydroponically controlled environments

Saplings of three riparian tree species (alder, birch and cottonwood) were grown for over 5 months in a hydroponics system that maintained the isotopic composition of source water in six treatments, ranging from –120 to +180‰δD and –15 to +10‰δ¹⁸O. The trees were grown in two greenhouses maintained at 25°C and at either 40 or 75% relative humidity, creating differences in transpiration rates and leaf water isotopic evaporative enrichment. The cellulose produced in the annual growth ring was linearly related to source water with differences in both slope and offset associated with greenhouse humidity. The slope of the isotopic composition of source water versus tree-ring cellulose was less than 1 for both δD and δ¹⁸O indicating incomplete isotopic exchange of carbohydrate substrate with xylem water during cellulose synthesis. Tests using the outer portion of the tree-ring and new roots were similar and showed that the tree-ring values were representative of the cellulose laid down under the imposed environmental conditions. The fraction of H and O in carbohydrate substrate that isotopically exchange with medium water was calculated to be 0.36 and 0.42 respectively, and biochemical mechanisms for these observed fractions are discussed. A mechanistic model of the biochemical fractionation events for both δD and δ¹⁸O leading to cellulose synthesis was robust over the wide range of cellulose stable isotope ratios. The experimental results indicate that both water source and humidity information are indeed recorded in tree-ring cellulose. These results help to resolve some of the disparate observations regarding the interpretation of stable isotope ratios in tree-rings found in the literature. Received: 4 January 1999 / Accepted: 12 August 1999     

A hierarchical analysis of minisatellite DNA diversity in Gambel oak (Quercus gambelii Nutt.; Fagaceae)

Gambel oak is a short shrub to medium-sized tree of southwest North America for which the potential to grow in large clonal stands has been proposed. Here we use three different multilocus VNTR DNA probes (synthesized via PCR) to demonstrate that clones growing together can be identified, but individual clones exceeding 50 m in diameter do not commonly occur in the study population of Quercus gambelii . Further, using a hierarchical sampling scheme (three circular transects with diameters of 9, 1100, and 28 000 m), we estimate that for nonclonal individuals: (i) mean number of bands analysed per individual = 22.92; (ii) mean similarity (band sharing) between individuals = 0.322; (iii) mean probability that two randomly chosen individuals share all bands = 4.93 × 10^–11; and (iv) mean estimated heterozygosity = 0.796. F _ST calculated for the two nonclonal levels of the sampling hierarchy was 0.023, indicating that little genetic differentiation exists between them. These results support previous findings that, due to life-history traits of the genus and this species (out-crossing, wind-pollinated, animal-dispersed, long-lived woody perennials), gene flow is high and genetic subdivision of populations is low in oaks. At this study site, the clonal nature demonstrated for Gambel oak appears to have little detectable effect on these population genetic characteristics.     

Plasticity in the degree of CAM-cycling and its relationship to drought stress in five species of Talinum (Portulacaceae)

Summary The degree of CAM-cycling was examined in plants from 23 populations representing five morphologically similar species of Talinum to determine how CAM-cycling correlates with site aridity and drought stress. In the field, CAM-cycling, as indicated by the amount of malic acid accumulated in plant tissues overnight, and stable carbon isotope ratio (¹³C) were positively correlated with an index of site aridity. The relative levels of CAM-cycling and ¹³C values among populations in the field reversed when plants were grown under less arid conditions in a growth chamber such that populations with the highest average CAM-cycling and ¹³C in the field had the lowest averages in the growth chamber. In both cases, plants from all populations showed significantly higher levels of CAM-cycling under drought-stressed conditions relative to conditions known or predicted to be less arid. CAM-cycling was also positively correlated with tissue water content in both well-watered and drought-stressed plants, possibly reflecting greater water conservation associated with reductions in stomatal conductance. Biomass accumulation in plants exhibiting the greatest degree of CAM-cycling in the growth chamber was suppressed by drought stress to a similar degree as in the other plants, yet reproductive biomass of these plants was inhibited to a smaller degree. Thus, the importance of CAM-cycling increases in these species of Talinum during drought stress, due to increased malic acid accumulation overnight, potentially enhancing their water status, survival, and reproduction.     

Purification and characterization of an anionic peroxidase from sycamore maple (Acer pseudoplatanus) cell suspension cultures

A 42 kDa anionic peroxidase (EC 1.11.1.7) having a pl of 3.6 was purified from suspension cultures of cells of sycamore maple ( Acer pseudoplatanus L.) grown in the dark by a combination of lectin-affinity, anion-exchange and gel permeation chromatography. The enzyme had an amino acid composition similar to that found for other anionic plant peroxidases, but the protein was blocked to amino-terminal protein sequencing. Commercially available antibodies against horseradish peroxidase were shown to cross-react with the sycamore maple enzyme on immunoblots. The purified peroxidase displayed differences in its affinity for each of the three monolignols, and these differences were compared to those found for a commercial preparation of horseradish peroxidase, as well as a laccase ( p -diphenol:O₂ oxidoreductase: EC 1.10.3.1) purified from sycamore maple cell suspension cultures. These results are discussed with respect to the role played by peroxidases in lignin deposition and host-pathogen response.     

Temporal patterns of inorganic nitrogen uptake by mature sugar maple (Acer saccharum Marsh.) and red spruce (Picea rubens Sarg.) trees using two common approaches

Background: Plant uptake of nitrogen influences many ecosystem processes, yet uptake by trees in northern forests of the United States has not been quantified throughout the growing season.

Aims: To measure NH₄ ⁺ and NO₃ ^? uptake by mature sugar maple (Acer saccharum) and red spruce (Picea rubens) trees during the early, mid and late growing season.

Methods: At Hubbard Brook Experimental Forest, New Hampshire, we used two approaches to measure nitrogen uptake capacity by mature trees: an in situ depletion method using intact roots and an ex situ ¹⁵N tracer method using excised roots.

Results: NH₄ ⁺ uptake was greater than NO₃ ^? for both methods and tree species (P < 0.05). NH₄ ⁺ uptake was lowest during the early growing season, while NO₃ ^? uptake was lowest during the late growing season. Measured rates of NH₄ ⁺ uptake were 2–3 orders of magnitude greater using the in situ depletion method compared with the ex situ ¹⁵N tracer method.

Conclusions: These results demonstrate seasonal differences in nitrogen uptake by two dominant tree species in a northern forest and show that the method employed can significantly impact measured rates of uptake, which could have implications for understanding the magnitude of plant nitrogen uptake and for cross-study comparisons of this process.     

Sexual reproductive biology of the endangered Japanese red maple (<Emphasis Type="Italic">Acer pycnanthum</Emphasis>)

Japanese red maple, Acer pycnanthum, is an endangered wetland species having dioecious sex expression. Prior studies conclude that land development has caused severe loss of habitat, and seedling regeneration is rare. However, information is lacking on sexual reproduction, which is the first critical stage of regeneration. My research objective was to describe flowering and fruiting characteristics of Japanese red maple, focusing on tree sizes at the onset of flowering, factors affecting flowering regularity and fruit abundance, periodicity of fruit production, and sex ratio. In a study of a young population, Japanese red maple initiated flowering at a small size under 100% light intensity. Initiation of flowering occurred at relatively lower height in males than females, and males flowered more regularly than females. A survey of 1,106 clones from 46 natural populations revealed that male clones were significantly more abundant than female clones. In a 4-year study of mature populations, nearly all clones flowered regularly, and fruit abundance of dominant females was typically high. Fruit abundance at the population level also remained high, although fruit abundance of individual female clones varied year by year. Therefore, sexual reproductive ability is highly vigorous. Sex ratio exhibited a major deviation from a 1:1 ratio in populations with a small number of clones. In conservation planning, we should prioritize avoidance of an unbalanced sex ratio by increasing population sizes. In addition, populations having many clones >40 cm dbh with large, well-lighted crowns have a high potential of regular and abundant seed production.     

Changes in water status and osmolyte contents in leaves and roots of olive plants (Olea europaea L.) subjected to water deficit

Two-year-old olive trees (Olea europaea L., cv. Coratina) were subjected to a 15-day period of water deficit, followed by 12 days of rewatering. Water deficit caused decreases in predawn leaf water potential (Ψ_w), relative water content and osmotic potential at full turgor (Ψ _π100) of leaves and roots, which were normally restored upon the subsequent rewatering. Extracts of leaves and roots of well-watered olive plants revealed that the most predominant sugars are mannitol and glucose, which account for more than 80% of non-structural carbohydrates and polyols. A marked increase in mannitol content occurred in tissues of water-stressed plants. During water deficit, the levels of glucose, sucrose and stachyose decreased in thin roots (with a diameter <1 mm), whereas medium roots (diameter of 1–5 mm) exhibited no differences. Inorganic cations largely contribute to Ψ _π100 and remained stable during the period of water deficit, except for the level of Ca²⁺, which increased of 25% in water-stressed plants. The amount of malate increased in both leaves and roots during the dry period, whereas citrate and oxalate decreased. Thin roots seem to be more sensitive to water deficit and its consequent effects, while medium roots present more reactivity and a higher osmotic adjustment. The results support the hypothesis that the observed decreases in Ψ_w and active osmotic adjustment in leaves and roots of water-stressed olive plants may be physiological responses to tolerate water deficit.     

Genotypic variation in drought response of silver birch (Betula pendula Roth): leaf and root morphology and carbon partitioning

This study investigates the drought response of four genotypes of Betula pendula with a focus on leaf and root morphological traits, leaf phenology and carbon partitioning between shoot and root. Potted one-year-old clonal plants of four genotypes from regions with low to high annual rainfall (550–1270 mm year⁻¹) were subjected to drought periods of 12–14 weeks in two subsequent years. Well-watered control plants of the four genotypes differed significantly with respect to total leaf area per plant (LA) and specific leaf area (SLA), whereas differences in total fine root surface area (RA), root specific area (SRA), and the fine root:leaf mass ratio (FR:LM) were not significant. Highest LA and SLA were found in the clone originating from the driest environment. In complementary physiological investigations this clone was found to have the highest water use as well which was interpreted as competitive superiority in terms of water consumption. Drought resulted in an increase in SLA in all genotypes, and a decrease in LA. Leaf area reduction was more pronounced in the genotypes from high than in those from low rainfall origin. The ratio of total root to leaf surfaces remained more or less constant after drought application despite an increase in FR:LM. This is explained by a decrease in SRA resulting from a reduced abundance of very small fine rootlets (diameter <0.2 mm) in the drought-treated plants. The loss in total root surface area due to a reduction in finest root mass was compensated for by a relative increase in total root dry mass per plant. Comparison of results from the first and second drought period indicated a marked influence of timing of drought, root system size, and putative root limitation on plant drought response. We conclude that leaf and root morphology, the total leaf and root surfaces, and the morphological response to drought in birch are to a large extent under genetic control.     

Accumulation of osmolytes and osmotic adjustment in water-stressed wheat (Triticum aestivum) and maize (Zea mays) as affected by calcium and its antagonists

Maize (Zea mays) and wheat (Triticum aestivum) were water stressed for 4 days at early vegetative growth (15-day-old) using PEG-6000 (−1.0 MPa), in the presence of 1 mM CaSO₄, 50 μM Verapamil (VP; calcium channel blocker); 50 μM Trifluoperazine (TFP; calmodulin antagonist) and then put to recovery in order to investigate the changes in osmoregulation in plants having C₃ and C₄ metabolism. Accumulation of proline (Pro) and quaternary ammonium compounds (QAC's), activities of pyrroline-5-carboxylate reductase (P5CR), proline dehydrogenase (PDH), water potential (Ψ_w), osmotic adjustment (OA), relative elongation rate (RER) and electrolyte leakage (EL) were examined during stress and recovery. Maize had significantly higher accumulation of Pro while wheat showed relatively more accumulation of QAC's. The activities of P5CR and PO were also significantly higher in maize than wheat. Maize shoots under stress showed higher Ψ_w, OA, RER and less EL than wheat shoots. Upon recovery from stress, maize regained its growth and water potential faster than wheat. Ca²⁺ elevated the accumulation of osmolytes in both the plants but OA was less sensitive to it. In the presence of Ca²⁺, wheat showed significantly more accumulation of osmolytes, higher Ψ_w, RER than maize. Ca²⁺ inhibitors partially reversed the effects of calcium indicating its involvement in governing solute accumulation. The differential sensitivity of maize and wheat towards water stress may be related to variation in endogenous calcium expression and its function.     

Position of mesquite (Prosopis spp) nodulation and nitrogen fixation (acetylene reduction) in 3-m long phraetophytically simulated soil columns

Summary The literature does not contain a field report of nodulation for the xerophytic tree legumes of the genus Prosopis despite their widespread occurrence in semi-arid regions of the world. A phraetophytically simulated greenhouse experiment was conducted with mesquite in a 3m deep soil column to determine if nodules would form in deper depths where moisture is more abundant. The upper 0.5m in the soil column was allowed to dry to 2200 kPa but the 3.2 m depth was maintained at soil moistures more positive than 70 kPa by water additions to the bottom of the soil column. Over 100 nodules and an acetylene reduction rate of 1.9 mg/h were observed at the 3.2m depth. Nodulation or acetylene reduction were not observed closer than 2.7m from the surface. Air temperatures during these assays exceeded 45°C. Leaf xylem water potentials were in the 2800–3500 kPa range.     

Differential use of large summer rainfall events by shrubs and grasses: a manipulative experiment in the Patagonian steppe

In the Patagonian steppe, years with above-average precipitation (wet years) are characterized by the occurrence of large rainfall events. The objective of this paper was to analyze the ability of shrubs and grasses to use these large events. Shrubs absorb water from the lower layers, grasses from the upper layers, intercepting water that would otherwise reach the layers exploited by shrubs. We hypothesized that both life-forms could use the large rainfalls and that the response of shrubs could be more affected by the presence of grasses than vice versa. We performed a field experiment using a factorial combination of water addition and life-form removal, and repeated it during the warm season of three successive years. The response variables were leaf growth, and soil and plant water potential. Grasses always responded to experimental large rainfall events, and their response was greater in dry than in wet years. Shrubs only used large rainfalls in the driest year, when the soil water potential in the deep layers was low. The presence or absence of one life-form did not modify the response of the other. The magnitude of the increase in soil water potential was much higher in dry than in humid years, suggesting an explanation for the differences among years in the magnitude of the response of shrubs and grasses. We propose that the generally reported poor response of deep-rooted shrubs to summer rainfalls could be because (1) the water is insufficient to reach deep soil layers, (2) the plants are in a dormant phenological status, and/or (3) deep soil layers have a high water potential. The two last situations may result in high deep-drainage losses, one of the most likely explanations for the elsewhere-reported low response of aboveground net primary production to precipitation during wet years. Received: 23 January 1997 / Accepted: 19 November 1997     

The influence of diet and water on the stable oxygen and hydrogen isotope composition of Chironomidae (Diptera) with paleoecological implications

Stable oxygen and hydrogen isotope analyses of fossil aquatic organisms, such as the chitinous head capsules of chironomid larvae (Chironomidae: Diptera), are promising proxies for inferring paleoecological conditions. In order for analyses of stable oxygen (δ¹⁸O) and hydrogen isotope ratios (δ²H) of fossil chironomid head capsules to be used effectively in paleoecological research, it is necessary to understand the factors controlling their stable oxygen and hydrogen composition. We cultured chironomid larvae in two isotopically distinct waters under controlled, replicated laboratory conditions. Chironomid larvae were fed on identical diets, to examine the degree to which water and diet influence the δ¹⁸O and δ²H of these organisms. We used a two-end member mixing model to determine the proportional contributions of oxygen and hydrogen from water to the oxygen and hydrogen of chironomid larvae. Our experiment demonstrated that 69.0 ± 0.4% of oxygen and 30.8 ± 2.6% of hydrogen in chironomid larvae are derived from habitat water. Our results show that oxygen isotopes from chironomid remains can better constrain past habitat water isotopic changes compared to hydrogen, due to 69% of the chironomid oxygen being influenced by habitat water. Our data add to a small but growing suite of comparative data on the sources of oxygen and hydrogen in animal tissues, and provide the first such analyses from aquatic insects.     

Geochemical study of vertebrate fossils from the Upper Cretaceous (Santonian) Csehbánya Formation (Hungary): Evidence for a freshwater habitat of mosasaurs and pycnodont fish

The diverse vertebrate remains from the Upper Cretaceous freshwater settings at Iharkút, Hungary, contain two fossil groups, Pycnodontiformes fish and Mosasauridae that are almost exclusively known from marine palaeo-environments. Hence, their appearance in alluvial sediments is very unusual. Trace element and isotope compositions of the remains have been analyzed to investigate the taphonomy and the ecological differences among the different fossil groups present at Iharkút.All examined fossils have undergone post-depositional diagenetic alteration, which resulted in high concentrations of REE, U, and Fe, together with almost complete homogenization of δ¹⁸O_CO3 values. Similar REE patterns in different fossils suggest a common origin for all remains, hence the discovered species most likely lived in the same local ecosystem. Despite partial diagenetic overprinting, the δ¹⁸O_PO4 values of the fossils indicate sufficient taxon-specific isotopic diversity to permit some broad conclusions on the palaeo-environment of the fossils. In particular, it is apparent that the isotopic composition of the Pycnodontiformes fish and Mosasauridae remains is most compatible with a freshwater palaeo-habitat and incompatible with a marine palaeo-environment. In addition, the Sr concentration and isotope data indicate that the Pycnodontiformes and Mosasauridae likely lived predominantly in a freshwater environment and were not simply occasional visitors to the Iharkút river ecosystem.Regarding other fossil groups, high δ¹⁸O_PO4 values of Alligatoroidea and Iharkutosuchus teeth suggest that these small crocodile species might have inhabited swamps and ponds where the water was relatively rich in ¹⁸O due to evaporation.     

Identification and characterization of phospholipase D and its association with drought susceptibilities in peanut (Arachis hypogaea)

Preharvest aflatoxin contamination has been identified by the peanut industry as a serious issue in food safety and human health because of the carcinogenic toxicity. Drought stress is the most important environmental factor exacerbating Aspergillus infection and aflatoxin contamination in peanut. The development of drought-tolerant peanut cultivars could reduce aflatoxin contamination and would represent a major advance in the peanut industry. In this study, we identified a novel PLD gene in peanut (Arachis hypogaea), encoding a putative phospholipase D (PLD, EC 3.1.4.4). The completed cDNA sequence was obtained by using the consensus-degenerated hybrid oligonucleotide primer strategy. The deduced amino acid sequence shows high identity with known PLDs, and has similar conserved domains. The PLD gene expression under drought stress has been studied using four peanut lines: Tifton 8 and A13 (both drought tolerant) and Georgia Green (moderate) and PI 196754 (drought sensitive). Northern analysis showed that PLD gene expression was induced faster by drought stress in the drought-sensitive lines than the drought tolerance lines. Southern analysis showed that cultivated peanut has multiple copies (3 to 5 copies) of the PLD gene. These results suggest that peanut PLD may be involved in drought sensitivity and tolerance responses. Peanut PLD gene expression may be useful as a tool in germplasm screening for drought tolerance. The nucleotide sequence, reported in this paper, have been submitted to GenBank under accession number AY274834.