Ecophysiological and morphological parameters related to survival in grass species exposed to an extreme climatic event

An experiment was performed to elucidate interspecific differences in survival time of grass species subjected to an extreme climatic event. We exposed eight grass species to a simulated heat wave in the field ('free air' temperature increase at 11°C above ambient) combined with drought. We determined whether interspecific differences in survival time were related to the responses of the species to the imposed stress or could be explained by their ecophysiological or morphological characteristics in unstressed conditions. Surprisingly, there was no effect of specific leaf area, but species with a higher total leaf area survived longer. This may arise from a greater water reserve in the plant as a whole, which could delay the desiccation of the meristem, or from reduced evaporation due to a higher leaf area index. Species in which the decrease in light-saturated stomatal conductance ( g _s ) and photosynthetic CO₂ uptake rate ( A _max ) was strongly related to the decrease in soil water availability (measured as soil relative water content and stress duration) survived longer than species in which g _s and A _max likewise declined but responded more to daily fluctuations in irradiance, temperature, and vapor pressure deficit during the heat wave. We, therefore, hypothesize that interspecific differences in stress survival time might be related to the extent to which stomata react to changes in soil water conditions relatively to changes in other environmental and physiological factors. The results suggest that resistance to extremes is governed by other mechanisms than resistance to moderate drought.     

Changes in body melanisation and desiccation resistance in highland vs. lowland populations of D. melanogaster

Wild caught samples of Drosophila melanogaster from five highland localities showed parallel changes in melanisation and desiccation resistance in darker versus lighter phenotypes, i.e. darker flies (>45% melanisation) showed significantly higher desiccation resistance than lighter flies (<30% melanisation). In order to find an association between body melanisation and desiccation resistance, highland and lowland populations from tropical and subtropical regions (11.15-31.06 degrees N) of the Indian subcontinent were raised and investigated at 21 degrees C for four physiological traits, i.e. per cent body melanisation, desiccation resistance, rate of water loss and rate of water absorption. On the basis of mother-offspring regression, body melanisation and desiccation resistance showed higher heritability (0.58-0.68) and thus these traits are suitable for laboratory analyses. Significantly higher melanisation as well as desiccation resistance were observed in highland populations as compared with lowland populations. The rates of water loss as well as absorption were negatively correlated with body melanisation, i.e. darker flies from highlands showed a reduced rate of water loss as well as a lower rate of water absorption while the reverse trend was observed in lighter flies from lowlands. On the basis of multiple regressions, significant effects due to combined altitude and latitude were observed for all the four physiological traits. Local climatic conditions (i.e. annual average temperature and relative humidity) helped in explaining parallel changes in body melanisation and desiccation resistance in D. melanogaster.     

Desiccation resistance of two semiterrestrial isopods, Ligia exotica and Ligia taiwanensis (Crustacea) in Taiwan

The ability to resist desiccation stress was examined in two semiterrestrial Ligia species, Ligia exotica Roux and L. taiwanensis Lee, in Taiwan, under a certain desiccation condition. L. exotica exhibited the longer survival time, lower weight-specific rates of water loss, and the slightly higher ability of tolerance to water loss, compared to L. taiwanensis. In each species, the animal size displays a positive correlation to the survival time and total ability to resist desiccation, yet this size effects on the weight-specific water loss rate is negative. Neither water content nor maximum tolerance to water loss shows the association with the animal size in both species. The path ways and magnitudes of the interactions between these traits of desiccation resistance are analyzed and diagrammed using a stepwise regression model. In this model, the body sizes of animal can explain the most part of the variations in the survival time. The body size has a direct effect and an indirect effect, through the effect on water loss rate, on the time that the experimental animals can survival under this desiccated condition. These results suggest that L. exotica attains larger size than does L. taiwanensis, a lower transpiration rate and, consequently, a greater ability in desiccation resistance. The performances of these interactions in the desiccated resistance are more advantageous for L. exotica to migrate and colonize in variable land habitats within a certain limit, and as a result that L. exotica shows a broader distribution pattern than did L. taiwanensis in Taiwan.     

Divergence of water balance mechanisms in two melanic Drosophila species from the western Himalayas

Drosophila busckii is more abundant under colder and drier montane habitats in the western Himalayas as compared to Drosophila melanogaster but the mechanistic basis of such climatic adaptations is largely unknown. We tested the hypothesis whether genetic variation or phenotypic plasticity of cuticular traits confer adaptive protection against desiccation stress in two melanic Drosophila species living under drier montane localities. For D. melanogaster, changes in melanisation are known to be associated with reduced water loss but there are no data on D. busckii. We investigated changes in body melanisation, cuticular lipids, desiccation resistance, water loss, extractable hemolymph volume (%), and dehydration tolerance in six sympatric populations of D. busckii and D. melanogaster over an altitudinal range of 640-2236 m. D. busckii is a melanic species but changes in cuticular water loss are negatively correlated with cuticular lipid mass and not with body melanisation. In D. melanogaster, there are no plastic effects (14-28 °C) for cuticular lipid mass but variation in body melanisation is associated with desiccation-related traits. Effects of organic solvents (hexane or chloroform: methanol), developmental plasticity and seasonal variation in cuticular lipids affect body water loss in D. busckii but no such changes occur in D. melanogaster. Thus, sympatric populations of D. busckii and D. melanogaster have evolved different water balance mechanisms under shared environmental conditions in the western Himalayas. Multiple measures of desiccation resistance in these species show clinal variation with altitude, consistent with adaptation to increased desiccation stress.     

Body melanization and its adaptive role in thermoregulation and tolerance against desiccating conditions in drosophilids

Melanism seems to have evolved independently through diverse mechanisms in various taxa and different ecological factors could be responsible for selective responses. Increased body melanization at higher altitudes as well as latitudes is generally considered to be adaptive for thermoregulation. Physiological traits such as body melanization and desiccation resistance have been investigated independently in diverse insect taxa at three levels: within populations, between populations and among species. A substantial number of Drosophila studies have reported clinal variations in both these traits along latitude. A possible link between these traits had remained unexplored in wild and laboratory populations of ectothermic insect taxa, including drosophilids, to date. Simultaneous analysis of these traits in assorted darker and lighter phenotypes in each population in the present study showed parallel changes for body melanization and desiccation resistance. The mechanistic basis of evolving desiccation resistance was explained on the basis of differential rates of water loss per hour in darker versus lighter phenotypes in six populations of Drosophila melanogaster from adjacent localities differing substantially in altitude all along the Indian subcontinent. Data on cuticular impermeability suggest a possible role of melanization in desiccation tolerance. However, substantial gaps remain in extending these results to other insect taxa and further exploring the physiological and molecular changes involved in melanization for conferring desiccation resistance.     

Revisiting water loss in insects: a large scale view

Desiccation resistance in insects has long been thought to covary with environmental water availability, and to involve changes in both cuticular and respiratory transpiration. Here, we adopt a large-scale approach to address both issues. Water loss rate and precipitation are positively related at global scales. A significant proportion (68%) of the interspecific variation in water loss rate is explained at the genus level or above. The relationship between metabolic rate and water loss rate differs substantially between mesic and xeric species. While these variables covary as a consequence of their independent covariation with body mass in mesic species, this is not the case in xeric species. In the latter, there is a strong relationship between the residuals of the water loss rate-body mass and metabolic rate-body mass relationships, and water loss rate is much reduced. Moreover, because metabolic rate does not differ significantly between xeric and mesic species of a similar size, respiratory transpiration constitutes a greater proportion of total water loss in xeric than in mesic species of a similar size. This implies that respiratory transpiration and the extent to which it can be modified must be of considerable importance in xeric insect species, although finer scale studies suggest otherwise.     

The importance of colony structure versus shoot morphology for the water balance of 22 subarctic bryophyte species

Questions: What are the water economy strategies of the dominant subarctic bryophytes in terms of colony and shoot traits? Can colony water retention capacity be predicted from morphological traits of both colonies and separate shoots? Are suites of water retention traits consistently related to bryophyte habitat and phylogenetic position? Location: Abisko Research Station, North Sweden. Methods: We screened 22 abundant subarctic bryophyte species from diverse habitats for water economy traits of shoots and colonies, including desiccation rates, water content at field capacity, volume and density (mg cm^?3) of water‐saturated and oven‐dried patches, evaporation rate (g·m^?2·s^?1) and cell wall thickness. The relationships between these traits and shoot and colony desiccation rates were analysed with Spearman rank correlations. Subsequent multivariate (cluster followed by PCA) analyses were based on turf density, turf and shoot desiccation rate, cell wall thickness and amount of external and internal water. Results: Individual shoot properties, i.e. leaf cell wall properties, water retention capacity and desiccation rate, did not correspond with colony water retention capacity. Colony desiccation rate depended on density of water‐saturated colonies, and was marginally significantly negatively correlated with species individual shoot desiccation rate but not related to any other shoot or colony trait. Multivariate analyses based on traits assumed to determine colony desiccation rate revealed six distinct species groups reflecting habitat choice and phylogenetic relationships. Conclusions: General relationships between shoot and colony traits as determinants of water economy will help to predict and upscale changes in hydrological function of bryophyte‐dominated peatlands undergoing climate‐induced shifts in species abundance, and feedbacks of such species shifts on permafrost insulation and carbon sequestration functions.     

Effects of age on water balance in Drosophila species.

Age-related declines in physiological performance have been documented in a wide variety of organisms. However, it is unknown whether related species age in a similar manner or whether physiological differences associated with aging differ widely among species. Previous work has shown that the desiccation resistance of Drosophila melanogaster decreases rapidly with age. Other Drosophila species may have delayed reproductive maturity or may inhabit arid habitats, so that age-related changes in water balance may be a significant physiological constraint on their ecological success. We may, therefore, predict that physiological systems involved in water balance will deteriorate with age at a reduced rate in these species. We quantified several components of water budgets for 14 species of Drosophila, including both desert and mesic representatives differing in their age of maturity. Desiccation resistance decreased with age in all but one species studied, primarily because of increased rates of water loss. However, there was no significant relationship between the rate of aging, as indicated by the rate of increase in water-loss rates as flies aged, and either habitat or age of maturity.     

Temperature and humidity acclimation increase desiccation resistance in the butterfly Bicyclus anynana

Desiccation resistance, that is, the ability to reduce water loss, is an ecologically important trait relevant to all terrestrial organisms, which may constrain species distributions. Nevertheless, relatively few studies have investigated plastic capacities in desiccation resistance. We here investigate plastic responses in body mass change, used as a proxy of desiccation resistance, to variation in temperature and relative humidity in the tropical butterfly Bicyclus anynana (Butler) (Lepidoptera: Nymphalidae). Our results indicate that butterflies acclimated to a higher (27 °C) compared with a lower temperature (18 °C) and a lower (50%) compared with a higher (90%) relative humidity displayed a decreased loss of body mass, and therefore likely a loss of body water (27 °C: 11%, 18 °C: 15%; 50% r.h.: 14%, 90% r.h.: 18%). Thus, mass loss was reduced under conditions indicating increased desiccation risk, suggesting adaptive phenotypic plasticity. Effects were most pronounced during the first 24 h after acclimation, indicating quick and transient responses to environmental conditions. As anthropogenic climate change is predicted to increase the magnitude and frequency of heat and drought periods, we argue that more studies on plastic capacities in traits relating to desiccation resistance are needed to better understand species responses.     

Divergent strategies for adaptation to desiccation stress in two Drosophila species of immigrans group

Water balance mechanisms have been investigated in desert Drosophila species of the subgenus Drosophila from North America, but changes in mesic species of subgenus Drosophila from other continents have received lesser attention. We found divergent strategies for coping with desiccation stress in two species of immigrans group--D. immigrans and D. nasuta. In contrast to clinal variation for body melanization in D. immigrans, cuticular lipid mass showed a positive cline in D. nasuta across a latitudinal transect (10°46'-31°43'N). Based on isofemale lines variability, body melanization showed positive correlation with desiccation resistance in D. immigrans but not in D. nasuta. The use of organic solvents has supported water proofing role of cuticular lipids in D. nasuta but not in D. immigrans. A comparative analysis of water budget of these two species showed that higher water content, reduced rate of water loss and greater dehydration tolerance confer higher desiccation resistance in D. immigrans while the reduced rate of water loss is the only possible mechanism to enhance desiccation tolerance in D. nasuta. We found that carbohydrates act as metabolic fuel during desiccation stress in both the species, whereas their rates of utilization differ significantly between these two species. Further, acclimation to dehydration stress improved desiccation resistance due to increase in the level of carbohydrates in D. immigrans but not in D. nasuta. Thus, populations of D. immigrans and D. nasuta have evolved different water balance mechanisms under shared environmental conditions. Multiple measures of desiccation resistance in D. immigrans but reduction in water loss in D. nasuta are consistent with their different levels of adaptive responses to wet and dry conditions on the Indian subcontinent.     

Invasive forbs differ functionally from native graminoids, but are similar to native forbs

? Exotic plant invasions can alter ecosystem processes, particularly if the invasive species are functionally different from native species. We investigated whether such alterations can be explained by differences in functional traits between native and invasive plants of the same functional group or by differences in functional group affiliation. ? We compared six invasive forbs in Europe with six native forbs and six native graminoids in leaf and whole-plant traits, plasticity in response to nutrient supply and interspecific competition, litter decomposition rate, effects on soil nutrient availability, and allelopathy. All traits were measured in a series of pot experiments, and leaf traits additionally in the field. ? Invasive forbs differed from native forbs for only a few traits; they had less leaf chlorophyll and lower phosphorus (P) uptake from soil, but they tended to have a stronger allelopathic effect. The invasive forbs differed in many traits from the native graminoids, their leaves had lower tissue densities and a shorter life span, their litter decomposed faster and they had a lower nitrogen-use efficiency. ? Our results suggest that invasive forbs have the potential to alter ecosystem properties when invading graminoid-dominated and displacing native graminoids but not when displacing native forbs.     

Trait convergence and plasticity among native and invasive species in resource-poor environments

? Premise of study: Functional trait comparisons provide a framework with which to assess invasion and invasion resistance. However, recent studies have found evidence for both trait convergence and divergence among coexisting dominant native and invasive species. Few studies have assessed how multiple stresses constrain trait values and plasticity, and no study has included direct measurements of nutrient conservation traits, which are critical to plants growing in low-resource environments. ? Methods: We evaluated how nutrient and water stresses affect growth and allocation, water potential and gas exchange, and nitrogen (N) allocation and use traits among a suite of six codominant species from the Intermountain West to determine trait values and plasticity. In the greenhouse, we grew our species under a full factorial combination of high and low N and water availability. We measured relative growth rate (RGR) and its components, total biomass, biomass allocation, midday water potential, photosynthetic rate, water-use efficiency (WUE), green leaf N, senesced leaf N, total N pools, N productivity, and photosynthetic N use efficiency. ? Key results: Overall, soil water availability constrained plant responses to N availability and was the major driver of plant trait variation in our analysis. Drought decreased plant biomass and RGR, limited N conservation, and led to increased WUE. For most traits, native and nonnative species were similarly plastic. ? Conclusions: Our data suggest native and invasive biomass dominants may converge on functionally similar traits and demonstrate comparable ability to respond to changes in resource availability.     

Divergent mechanisms for water conservation in Drosophila species

The role of melanization and cuticular lipids in water conservation has been studied in many Drosophila species (Diptera: Drosophilidae). Nevertheless, a comparative approach to larval and adult stages of ecologically diverse, wild Drosophila species is still required. Based upon abdominal cuticular melanization patterns, wild‐caught Drosophila species were categorized as (1) melanic, (2) fixed‐melanic, or (3) non‐melanic. At the interspecific level, the ecological significance of melanization and cuticular lipids was determined by the inverse association of melanization and cuticular water loss in melanic species, and of cuticular lipids and cuticular water loss in fixed‐melanic and non‐melanic species. Interestingly, higher amounts of cuticular lipids were also evident in fixed as well as non‐melanic species, as compared to melanic species at larval stages, which is consistent with their differences in reduced water loss rates. Moreover, fixed‐melanic and non‐melanic species exhibited comparatively higher (ca. 1.8–2.0 fold) desiccation resistance. Thus, cuticular lipids provide a better waterproofing mechanism than melanization. Furthermore, acclimation to dehydration stress in adults improved desiccation resistance in melanic species, whereas such effects were lacking in fixed‐melanic and non‐melanic species. However, there were no changes in cuticular components as a consequence of desiccation acclimation. Thus, our results indicate that melanic, fixed‐melanic, and non‐melanic Drosophila species differ in the evolved physiological mechanisms of water conservation to adapt to dry conditions.     

Relationship between species relative abundance and plant traits for an infertile habitat

This study tested whether differences in species abundance at an infertile site could be explained by differences in the species' plant traits. Nine traits were chosen for the analysis based on results of previous studies conducted across soil fertility gradients. The traits were measured for each of seven herbaceous species whose abundance ranged from 5% to 100% of locations occupied in a ridgetop habitat. Using linear regression, significant relationships were found between species relative abundance and each of five traits. In these relationships, a trait explained between 69% and 88% of interspecific variation in abundance. Relatively abundant species had a slower growth rate, smaller shoot mass, higher root to shoot ratio, slower loss of leaf tissue to herbivores and higher infection of roots by mycorrhizal fungi than less abundant species. Using three of these five traits (i.e. shoot mass, mycorrhizal infection and loss of leaf tissue to herbivores) as independent variables in a multiple regression equation explained 99% of interspecific variation in abundance. The latter result indicates that species relative abundance can be explained for a single habitat by choosing traits found to be related to species abundance in previous gradient studies. However, not every trait chosen was significantly related to species abundance. Therefore, a large number of traits may have to be chosen initially to ensure that some subset of these traits can explain species relative abundance.     

Growth of tree seedlings in a tropical dry forest in relation to soil moisture and leaf traits

"AimsThe growth of plant species in tropical dry forest (TDF) is expected to be largely governed by the availability of soil moisture. In this study we attempt to identify mechanisms by which seedlings of dry tropical trees cope with water stress by adjusting their leaf characteristics to water availability and micro environments, and address following questions: How are leaf traits and relative growth rate (RGR) of the dominant seedling species of TDF affected by seasonal changes in soil moisture content (SMC)? What is the relationship of functional traits with each other? Can leaf traits singly or in combination predict the growth rate of seedling species of TDF? The study was conducted in situ on four sites (viz., Hathinala, Gaighat, Harnakachar and Ranitali, listed in order of decreasing SMC) within the tropical dry deciduous forest in northern India. Methods Five leaf traits viz., specific leaf area (SLA), leaf dry matter content (LDMC), concentrations of leaf nitrogen (leaf N), phosphorus (leaf P) and chlorophyll (Chl) and two physiological processes, viz., stomatal conductance (Gs net) and photosynthetic rate (A net), and RGR, of four dominant tree seedling species of a TDF (viz., Buchanania lanzan, Diospyros melanoxylon, Shorea robusta and Terminalia tomentosa) on four sites were analysed for species, site and season effects over a 2-year period. Step-wise multiple regression was performed to predict RGR from mean values of SMC, leaf traits and physiological processes. Principal component analysis (PCA) was performed to observe the extent of intra- vs. inter-specific variability in the leaf traits and physiological rates.Important findings All the traits and physiological rates were interrelated and showed significant positive relationship with RGR except for the correlation of LDMC with RGR which was not significant. Further, relationships of SMC with all leaf traits, physiological rates and RGR were significant, except for that between SMC and SLA for B. lanzan and D. melanoxylon. The slope of seedling trait:SMC relationship, a measure of phenotypic plasticity in response to soil moisture gradient, varied among species. Among the four species, T. tomentosa was the most plastic and S. robusta the least. In conclusion, leaf traits and physiological processes were strongly related to soil water availability on the one hand and seedling growth on the other. Gs net is the most important variable which accounted for the greatest amount of variability (62%) in RGR, emphasizing the role of stomatal conductance in shaping growth patterns across spatial and temporal gradients of soil water availability. Gs net and SMC together explained 64% variability in RGR, indicating that other traits/factors, not studied by us are also important in modulating the growth of tropical tree seedlings.     

Developmental and acclimatory contributions to water loss in a desert rodent: investigating the time course of adaptive change

Understanding the evolution of physiological traits requires considering three nonexclusive mechanisms that underlie phenotypes and cause their change over different time scales: acclimation, developmental plasticity, and natural selection for genetically fixed traits. Physiological adjustments to changes in the desiccating potential of the environment were investigated with one subspecies of common desert rodent, Dipodomys merriami merriami (Merriam's kangaroo rat). We raised young whose parents originated from environments that differ in both temperature and humidity. These young were raised under either desiccating or water-abundant conditions, and their water loss was measured at a series of temperatures to determine the effect developmental conditions have on resistance to desiccation. We then determined the contribution of acclimation to desiccation resistance by keeping the differentially raised young in conditions opposite to those during their development and again measuring water loss. We found that developmental plasticity and acclimation can completely account for the existing intraspecific variability in desiccation resistance under certain conditions. In fact, developmental and acclimatory changes can equal genetically based differences of the populations. This phenotypic plasticity can operate relatively quickly and therefore may attenuate the actions of natural selection. Understanding the extent and nature of such flexibility is critical to our understanding intraspecific variability and the consequences of changing climate.     

Geographic variation for climatic stress resistance traits in the springtail Orchesella cincta

Multiple traits of stress resistance were investigated in the epedaphic springtail Orchesella cincta. Second generation adults from five laboratory populations were compared with respect to resistance to extreme temperatures and desiccation, and traits relevant to climatic adaptation. Populations were collected along a 2000-km latitudinal gradient ranging from Denmark to southern Italy and reared under the same standard laboratory conditions. Traits investigated were resistance to high and low temperature, desiccation resistance, body size and water loss rate (WLR). Results showed genetically based differences in resistance to high and low temperature, desiccation, WLR, water pool and body size between populations. Individuals from the most northern population had the highest desiccation-and cold shock resistance, and the lowest heat shock resistance. Females were significantly more desiccation resistant than males. The results of cold shock resistance showed a positive increase with lowest environmental temperature recorded at the sites of population origin, whereas heat shock resistance showed a positive increase with highest recorded temperature at the sites of population origin. Desiccation resistance increased towards the most southern and northern population, suggesting that both low and high temperature extremes affect desiccation resistance. Body mass, water pool and WLR showed interpopulation as well as sex specific variation. This provides evidence for geographical variation in stress resistance of springtails related to climatic conditions.     

Relation between Water Balance and Climatic Variables Associated with the Geographical Distribution of Anurans

Amphibian species richness increases toward the equator, particularly in humid tropical forests. This relation between amphibian species richness and environmental water availability has been proposed to be a consequence of their high rates of evaporative water loss. In this way, traits that estimate water balance are expected to covary with climate and constrain a species’ geographic distribution. Furthermore, we predicted that coexisting species of anurans would have traits that are adapted to local hydric conditions. We compared the traits that describe water balance in 17 species of anurans that occur in the mesic Atlantic Forest and xeric Cerrado (savannah) habitats of Brazil. We predicted that species found in the warmer and dryer areas would show a lower sensitivity of locomotor performance to dehydration (SLPD), increased resistance to evaporative water loss (REWL) and higher rates of water uptake (RWU) than species restricted to the more mesic areas. We estimated the allometric relations between the hydric traits and body mass using phylogenetic generalized least squares. These regressions showed that REWL scaled negatively with body mass, whereas RWU scaled positively with body mass. Additionally, species inhabiting areas characterized by higher and more seasonally uniform temperatures, and lower and more seasonally concentrated precipitation, such as the Cerrado, had higher RWU and SLPD than species with geographical distributions more restricted to mesic environments, such as the Atlantic Forest. These results support the hypothesis that the interspecific variation of physiological traits shows an adaptation pattern to abiotic environmental traits.     

Seasonal changes in humidity level in the tropics impact body color polymorphism and desiccation resistance in Drosophila jambulina—Evidence for melanism-desiccation hypothesis

Drosophila jambulina exhibits color dimorphism controlled by a single locus but its ecological significance is not clear. Dark and light morphs differ significantly in body melanisation, desiccation resistance, rate of water loss, mating activity and fecundity. Interestingly, this species lacks clinal variation for body size, desiccation resistance and life history traits. For body melanisation, lack of geographical variation as well as plastic effects is not in agreement with a thermal melanism hypothesis. However, based on field data, there are seasonal changes in phenotypic frequencies of dark and light body color morphs which correlate significantly with variation in humidity levels. Under short-term (8 h) desiccation stress, we observed higher number of assortative matings, longer copulation period and increased fecundity for dark strains as compared with light strains. By contrast, both the morphs when exposed to high humid conditions exhibited higher assortative matings and fecundity for light strains as compared with dark strains. In tropical populations of D. jambulina, body color polymorphism seems to be maintained through humidity changes as opposed to thermal melanism. Thus, seasonal changes in the frequency of body color morphs in this tropical species supports melanism-desiccation hypothesis.