Genetic Variation of Drought Tolerance in Pinus pinaster at Three Hierarchical Levels: A Comparison of Induced Osmotic Stress and Field Testing

Understanding the survival capacity of forest trees to periods of severe water stress could improve knowledge of the adaptive potential of different species under future climatic scenarios. In long lived organisms, like forest trees, the combination of induced osmotic stress treatments and field testing can elucidate the role of drought tolerance during the early stages of establishment, the most critical in the life of the species. We performed a Polyethylene glycol-osmotic induced stress experiment and evaluated two common garden experiments (xeric and mesic sites) to test for survival and growth of a wide range clonal collection of Maritime pine. This study demonstrates the importance of additive vs non additive effects for drought tolerance traits in Pinus pinaster, and shows differences in parameters determining the adaptive trajectories of populations and family and clones within populations. The results show that osmotic adjustment plays an important role in population variation, while biomass allocation and hydric content greatly influence survival at population level. Survival in the induced osmotic stress experiment presented significant correlations with survival in the xeric site, and height growth at the mesic site, at population level, indicating constraints of adaptation for those traits, while at the within population level no significant correlation existed. These results demonstrate that population differentiation and within population genetic variation for drought tolerance follow different patterns.     

Heritable variation in the survival of seedlings from Patagonian cypress marginal xeric populations coping with drought and extreme cold

The rear edges of tree species have begun to be perceived as highly valuable for genetic resources conservation and management. In view of expected climatic changes, the responses of trees at their xeric limits may largely be determined by their capacity to cope with augmented environmental variance. We assess the heritability of early survival of Patagonian cypress in two common-garden field tests with contrasting summer water deficits, comprising 140 and 163 open-pollinated families from 10 marginal xeric populations. The first experiment underwent less rigorous conditions than the average mesic, Mediterranean climatic conditions, which were sufficient to reveal additive genetic effects of summer drought on seedling survival. The second trial suffered strong summer water-deficit stress and a winter extreme cold event. In this harsher environment, the heritabilities of survival under summer water-deficit stress were high in all the populations (h ²?=?0.84 on average), while the heritabilities of seasonal, extreme cold survival were moderate or even nil (h ²?=?0.28 on average). We did not find evidence of genetic differentiation among populations in their capabilities to survive droughts and cold extremes. Our results indicate that even when climatic changes were strong enough to cause the extinction of the most threatened populations, heritable variation for traits underlying drought and cold tolerances may allow the marginal xeric edge of cypress to persist under augmented environmental variance, without losing overall genetic diversity.     

Gender-Related developmental instability and herbivory ofPistacia atlantica across a steep environmental gradient

To examine gender-related developmental instability and herbivory of a plant across a steep climatic gradient, we studiedPistacia atlantica DESF., a dioecious tree of the Irano-Turanian zone in the Middle East. Leaves were sampled from five populations along a climatic gradient in Israel from mesic Mediterranean to xeric (and geographically peripheral) sites. Leaves of the xeric populations were ca. 25% smaller in size (length) than leaves of other populations. Leaf developmental instability (an indicator of stress) was examined using the measurement of leaf asymmetry (LA), estimated by the deviation from the origin of paired lateral leaflets and by the presence vs. absence of the terminal leaflet. LA was not correlated with the climatic (site) gradient and had the highest, though not significantly, levels in the intermediate ecotonal zone between the Mediterranean and the arid ecosystems. Male trees were 13% higher in LA than females. Density of two specialist gall-forming aphids varied with site but was not affected by plant gender. Gender-related developmental instability is discussed in relation to resource allocation in early spring. Because the study was limited to five populations further research is clearly needed.     

The Effect of Age, Pre-conditioning, and Water Stress on the Transpiration Rates of Douglas-Fir (Pseudotsuga menziesii) Seedlings of Several Ecotypes

Seedlings of Douglas-fir from seed of a number of mesic and xeric origins were grown in growth chambers and a nursery to various ages up to 16 weeks. Measurements were made to determine the effect of seedling age, growth chamber and nursery pre-conditioning, and seed source on transpiration rates under closely controlled laboratory conditions. Additional experiments were conducted on seedlings of two contrasting ecotypes to determine the effect of different pre-conditioning combinations of plant and soil water potential on seedling transpiration rates. Results show that well-watered seedlings of two mesic ecotypes show no decline in transpiration rates per unit leaf area up to 16 weeks of age while corresponding seedlings of three exeric ecotypes do decline. The growth chamber pre-conditioning results in lower seedling transpiration rates and more decline in seedling transpiration rates with increasing plant water stress than for nursery pre-conditioning. In a similar way, the xeric ecotype seedlings have more decline in transpiration rates with increasing plant water stress than do the mesic ecotype seedlings. Soil water potential influences transpiration rates through pre-conditioning effects. Seedlings which have experienced prior soil moisture stress decrease transpiration more in response to low plant water potentials than do plants which have experienced no soil moisture stress. These behavioral characteristics illustrate adaptive means by which seedlings conserve water through the interaction of genetic and preconditioning mechanisms.     

Contrasting growth changes in two dominant species of a Mediterranean shrubland submitted to experimental drought and warming

BACKGROUND AND AIMS: Climate projections predict drier and warmer conditions in the Mediterranean basin in the next decades. The possibility of such climatic changes modifying the growth of two Mediterranean species, Erica multiflora and Globularia alypum, which are common components of Mediterranean shrublands, was assessed. METHODS: A field experiment was performed from March 1999 to March 2002 to prolong the drought period and to increase the night-time temperature in a Mediterranean shrubland, where E. multiflora and G. alypum are the dominant species. Annual growth in stem diameter and length of both species was measured and annual stem biomass production was estimated for 1999, 2000 and 2001. Plant seasonal growth was also assessed. KEY RESULTS: On average, drought treatment reduced soil moisture 22 %, and warming increased temperature by 0.7-1.6 degrees C. Erica multiflora plants in the drought treatment showed a 46 % lower annual stem elongation than controls. The decrease in water availability also reduced by 31 % the annual stem diameter increment and by 43 % the annual stem elongation of G. alypum plants. New shoot growth of G. alypum was also strongly reduced. Allometrically estimated biomass production was decreased by drought in both species. Warming treatment produced contrasting effects on the growth patterns of these species. Warmer conditions increased, on average, the stem basal diameter growth of E. multiflora plants by 35 %, raising also their estimated stem biomass production. On the contrary, plants of G. alypum in the warming treatment showed a 14 % lower annual stem growth in basal diameter and shorter new shoots in spring compared with controls. CONCLUSIONS: The results indicate changes in the annual productivity of these Mediterranean shrubs under near future drier and warmer conditions. They also point to alterations in their competitive abilities, which could lead to changes in the species composition of these ecosystems in the long term.     

Maternal effects of drought stress and inbreeding in Impatiens capensis (Balsaminaceae)

Maternal effects can have substantial impacts on plant fitness and plant populations. Stressful environmental conditions can cause a maternal plant to inadequately provision its progeny, resulting in poor seedling growth, low reproductive success, and decreased competitive ability. Maternal effects consist of environmental and genetic load components, but the interactions between these two components have rarely been considered. To determine the effects of maternal drought stress and maternal inbreeding on progeny biomass (a fitness correlate) and physiological responses to drought stress, we conducted a greenhouse experiment with genetic lines from two populations (mesic site vs. dry site) of the herbaceous annual Impatiens capensis (Balsaminaceae). Seeds were collected from cleistogamous flowers of inbred or outcrossed maternal plants that were subject to either a drought or control treatment. These seeds were grown into juvenile plants that were also subject to either a drought stress or a control treatment. Plants from the mesic site had significantly reduced biomass from maternal drought stress, while plants from the dry site maintained biomass despite adverse maternal environmental conditions. Juvenile plants of both populations had reduced biomass only as a result of maternal inbreeding. Interestingly, inbreeding depression was more apparent when maternal environmental conditions were benign.     

Patterns of functional clonal traits and clonal growth modes in contrasting grasslands in the central Apennines,Italy

Abstract. Aim: Patterns of plant functional traits related to clonality (clonal growth modes; CGM) in plant communities were studied and hypotheses on the importance of the selected traits in plant communities supported by soils differing in moisture and nutrient status were tested. Material and Methods: Selected plant functional traits, such as the position of the mother‐daughter plants connections, length of spacers, frequency of multiplication, persistency of ramets connections, presence of storage organs and bud protection were studied in two contrasting plant communities (xeric and mesic abandoned pastures) typical of central Apennines, Italy. Results and Discussion: Clonality was shown to be of great importance in both mesic and xeric grasslands. The major differences between the two communities were due to the dominant CGMs: turf graminoids (having effective protection of growth meristems in dense tussocks) dominated xeric grasslands, while rhizomatous graminoids (typical of competitive resource‐rich environments) dominated mesic grasslands. Below‐ground CGOs (clonal growth organs), shorter spacers, higher multiplication potential, permanent ramet connection, large bud bank and increased importance of bud protection were found to be of importance in water stressed xeric grassland. Contrary to our expectations, the mesic (less stressed) grasslands have the higher number of clonal plants possessing storage organs.     

Twig morphology and anatomy of Mediterranean trees and shrubs related to drought tolerance

De Micco V. and Aronne G. 2008. Twig morphology and anatomy of Mediterranean trees and shrubs related to drought tolerance. Bot. Helv. 118: 139 – 148. Woody species populating Mediterranean ecosystems have evolved phenological, physiological and morphological adaptations to summer drought, but their degree of drought tolerance varies, so that different species dominate along gradients from xeric to more mesic environments. In this study, we analysed morpho-anatomical properties of leaves and twigs of eight Mediterranean woody species. To test whether there is a consistent pattern of adaptation to drought, we hypothesised that the rank order of species along the xeric-mesic gradient would be reflected by increasing or decreasing trends in twig properties or in relationships between leaf area and branch size. One-year-old twigs were sampled from plants growing in a common site on the Tresino promontory in southern Italy. Measurements included leaf number and area, branch length and diameter, as well as the relative amounts of pith, xylem and cortex tissues at the basis of branches. All traits varied significantly among species, but none of them correlated with the rank order of species along the xeric-mesic gradient. In addition, the relationship between leaf area and branch size (length or diameter) was not consistently related to the drought tolerance of the species. Inconsistencies could partly be explained by variations in twig anatomy. These results support the view that not single traits, but combinations of morpho-anatomical features influence the overall capability of species to withstand dry conditions. Submitted 12 May 2008; Accepted 24 September 2008 Subject editor: Sabine Güsewell     

Life history variation in an annual plant under two opposing environmental constraints along an aridity gradient

Environmental gradients represent an ideal framework for studying adaptive variation in the life history of plant species. However, on very steep gradients, largely contrasting conditions at the two gradient ends often limit the distribution of the same species across the whole range of environmental conditions. Here, we study phenotypic variation in a winter annual crucifer Biscutella didyma persisting along a steep gradient of increasing rainfall in Israel. In particular, we explored whether the life history at the arid end of the gradient indicates adaptations to drought and unpredictable conditions, while adaptations to the highly competitive environment prevail at the mesic Mediterranean end. We examined several morphological and reproductive traits in four natural populations and in populations cultivated in standard common environment. Plants from arid environments were faster in phenological development, more branched in architecture and tended to maximize reproduction, while the Mediterranean plants invested mainly in vertical vegetative growth. Differences between cultivation and field in diaspore production were very large for arid populations as opposed to Mediterranean ones, indicating a larger potential to increase reproduction under favorable conditions. Our overall findings indicate two strongly opposing selective forces at the two extremes of the aridity gradient, which result in contrasting strategies within the studied annual plant species.     

Radial-growth and wood-anatomical changes in overaged Quercus pyrenaica coppice stands: functional responses in a new Mediterranean landscape

Recent land-use changes in intensively managed forests such as Mediterranean coppice stands might profoundly alter their structure and function. We assessed how the abandonment of traditional management practices in coppice stands, which consisted of short cutting-cycles (10–15 years), has caused overaging (stems are usually much older than when they were coppiced) and altered their wood anatomy and hydraulic architecture. We studied the recent changes of wood anatomy, radial growth, and hydraulic architecture in two stands of Quercus pyrenaica, a transitional Mediterranean oak with ring-porous wood forming coppice stands in W–NW Spain. We selected a xeric and a mesic site because of their contrasting climates and disturbance histories. The xeric site experienced an intense defoliation after the severe 1993–1994 summer drought. The mesic site was thinned in late 1994. We studied the temporal variability in width, vessel number and diameter, and predicted the hydraulic conductivities (K _h) of earlywood and latewood. In the mesic site, we estimated the vulnerability to xylem cavitation of earlywood vessels. Overaging caused a steep decline in latewood production at a cambial age of 14 years., which was close to the customary cutting cycle of Q. pyrenaica. The diameter distribution of vessels was bimodal, and latewood vessels only accounted for 4% of the K _h. Overaging, acting as a predisposing factor in the decline episode, was observed at the xeric site, where most trees did not produce latewood in 1993–1995. At the mesic site, thinned trees formed wider tree-rings, more latewood and multiseriate tree-rings than overaged trees. The growth enhancement remained 8 years after thinning. Most of the hydraulic conductivity in earlywood was lost in a narrow range of potentials, between −2.5 and −3.5 MPa. We have shown how hydraulic conductivity and radial growth are closely related in Q. pyrenaica and how aging modulates this relationship.     

Physiological performance of silver-fir (Abies alba Mill.) populations under contrasting climates near the south-western distribution limit of the species

Silver-fir (Abies alba Mill.) populations located at the south side of the main Pyrenean axis and Pre-Pyrenees constitute the south-western distribution limit of the species and, therefore, may be responding more noticeably to climate change than those in the centre of the range. The increasing aridity detected in the Pyrenees during the 20th century should affect more negatively the physiological performance of the southernmost silver-fir stand growing under more xeric conditions in comparison with stands growing within the main distribution area under mesic conditions. To evaluate the climatic influence on the performance of silver fir near its distribution limit, we studied several physiological and growth variables in shoots and needles from two silver fir populations located in nearby but climatically contrasting sites: Paco Ezpela (site E) and Gamueta (site G). Site E showed a stronger Mediterranean influence than site G, i.e. the former site was characterized by higher temperatures and leaf-to-air water vapour pressure difference and lower precipitation in summer than the latter site. Silver firs from site E showed lower values of primary and secondary growth, needle length, stomatal conductance, net photosynthesis and photosystem II (PSII) efficiency than individuals from site G. The reduction in net photosynthesis could be ascribed to a low CO₂ availability and to a lower PSII efficiency. We conclude that the physiological differences found between both sites were caused by the more xeric conditions of site E as compared with the more mesic environment in site G. The predicted increase of severe droughts in the southern Pyrenees might cause a decrease in photosynthesis and growth in those silver-fir stands located near the ecological limit of the species.     

Living on the edge: Legacy of water availability on Tetraclinis articulata secondary growth under semiarid conditions in Morocco

Tetraclinis articulata is a xerothermic Mediterranean conifer native from NW Africa that grows under semiarid conditions in the Atlas Mountains. This species is particularly well adapted to water scarcity and forms xeric woodlands subjected to recurring drought, aridification and overexploitation. Wood of T. articulata has been considered of limited dendrochronological value due to abundant anatomical anomalies in their growth rings. We studied tree-ring growth and tree-ring features to check its dendrochronological potential and evaluate their climatic signals and drought legacy effects at two semiarid sites with contrasting elevation in Morocco. Tree-ring boundaries were properly identified on wood cores from 60 to 68% of the sampled trees. Intra-annual wood density fluctuations (IADF), microrings, missing rings and rings with undefined limits were most abundant at the low-elevation coastal site than at the mountain site. Microring and missing ring abundance was inversely related to tree-ring growth, independent of cambial age, and mostly dependent on drought occurrence. IADF frequency was independent of tree-ring growth but inversely related to cambial age. A complex response of IADF formation to cool March to June conditions was found at the low-elevation coastal site, while IADF formation in the mountain site was related to May water availability. Undefined ring limits were observed only at the low-elevation site, showing positive relationships with elevated maximum temperatures in previous winter. Previous winter rainfall favored radial growth, but a significant legacy effect of water availability was evident up to three years prior to growth. Tree-ring growth was mostly related to water availability 33 and 26 months prior to growth at low-elevation and mountain sites, respectively. The obtained results reveal a very plastic cambial activity and a strong ability of T. articulata to withstand drought even for long periods.     

Tests of the contribution of acclimation to geographic variation in water loss rates of the West Indian lizard <Emphasis Type="Italic">Anolis cristatellus</Emphasis>

Phenotypic plasticity can contribute to the process of adaptive radiation by facilitating population persistence in novel environments. West Indian Anolis lizards provide a classic example of an adaptive radiation, in which divergence has occurred along two primary ecological axes: structural microhabitat and climate. Adaptive plasticity in limb morphology is hypothesized to have facilitated divergence along the structural niche axis in Anolis, but very little work has explored plasticity in physiological traits. Here, we experimentally ask whether Puerto Rican Anolis cristatellus from mesic and xeric habitats differ in desiccation rates, and whether these lizards exhibit an acclimation response to changes in relative humidity. We first present microclimatic data collected at lizard perch sites that demonstrate that abiotic conditions experienced by lizards differ between mesic and xeric habitat types. In Experiment 1, we measured desiccation rates of lizards from both habitats maintained under identical laboratory conditions. This experiment demonstrated that desiccation rates differ between populations; xeric lizards lose water more slowly than mesic lizards. In Experiment 2, lizards from each habitat were either maintained under the conditions of Experiment 1, or under extremely low relative humidity. Desiccation rates did not differ between lizards from the same habitat maintained under different treatments and xeric lizards maintained lower desiccation rates than mesic lizards within each treatment. Our results demonstrate that A. cristatellus does not exhibit an acclimation response to abrupt changes of hydric conditions, and suggest that tropical Anolis lizards might be unable to exhibit physiological plasticity in desiccation rates in response to varying climatic conditions.     

Phenology constrains opportunistic growth response in <Emphasis Type="Italic">Bromus tectorum</Emphasis> L.

Seasonal resource availability may act as a constraint on plant phenology and thereby influence the range of growth responses observed among populations of annual species, especially those occupying a wide range of environments. We compared a mesic and a xeric population of the non-native, annual grass, Bromus tectorum, to examine phenology in response to interspecific competition and water availability. Using a target-neighborhood approach, we assessed how phenological patterns of the two populations affected morphological and growth responses to enhanced resource availability represented by late-season soil moisture. The xeric population exhibited a highly constrained phenology and was unable to extend the growing season despite available soil resources. Because of the low phenotypic variation, allocation to reproduction was similar across resource conditions. In contrast, the mesic population flowered later and showed a more opportunistic phenology in response to late-season water availability. The mesic population was not able to maintain consistent reproductive allocation at low resource levels. The responses of the two populations to late-season water availability were not affected by the density of neighboring plants. We suggest that post-introduction selection pressure on B. tectorum in the xeric habitat has resulted in a more fixed phenology which limits opportunistic response to unpredictable, particularly late-season resource availability. Opportunistic and fixed responses represent contrasting strategies for optimizing fitness in temporally varying environments and, while both play important roles for ensuring reproductive success, these results suggest that local adaptation to temporal resource variation may reflect a balance between flexible and inflexible phenology.     

Photoprotection mechanisms in Quercus ilex under contrasting climatic conditions

Acclimation to local conditions may produce adaptive responses in plants subjected to diverse climatic stresses. However, it has not been assessed how local adaptation and phenotypic plasticity contribute to photoprotection mechanisms in response to contrasting climatic conditions in Mediterranean tree species.     

Balance of Competitive and Facilitative Effects of Exotic Trees on a Native Patagonian Grass

The balance between facilitation and competition in plants changes with species characteristics and environmental conditions. Facilitative effects are common in natural ecosystems, particularly in stressful environments or years. Contrarily, in artificial associations of plants, such as agroforestry systems, some authors have suggested that even when facilitative effects may occur, net balance of tree effects on grasses is usually negative, particularly in dry environments. The aim of this study was to determine the net effect of the exotic ponderosa pine on the native grass Festuca pallescens (St. Ives) Parodi in agroforestry systems in Patagonia. Soil water content, plant water status, and relative growth were measured in the grass growing in different treatments (determined by tree cover level) during two growing seasons with contrasting climatic conditions. Facilitative effects of trees over grass water status were recorded only when water availability was high. A net negative effect was detected on dates when soil water content was very low and evaporative demand was high. The strength of these negative effects depended on tree density and climatic conditions, being higher in treatments with lower tree canopy cover. These results indicate that the positive effect of trees could only be expected under relatively low stress conditions. However, relative growth of grasses was always similar in plants growing in forested plots than in open grassland. Differences in biomass allocation for grasses growing in shade and open habitats may reconcile these contrary results. Our results highlight the importance of the physiology of a species (relative drought and shade tolerance) in determining the response of a plant to a particular interacting species.     

Spring water deficit and soil conditions matter more than seed origin and summer drought for the establishment of temperate conifers

In anticipation of more severe summer droughts, forestry in temperate Europe is searching for drought-resistant ecotypes of native tree species that might maintain ecosystem services in the future. We investigated how spring precipitation and soil conditions interact with summer drought and affect the establishment of conifer seedlings from different climatic origin. Emergence, establishment and subsequent performance of seedlings originating from autochthonous, Central Alpine, continental Eastern European, and Mediterranean Pinus sylvestris and Picea abies populations were studied in the dry Alpine Rhine valley, Switzerland, at three sites with differing soil water holding capacities and in 3 years with contrasting weather conditions. In addition to this natural inter-annual variation, precipitation was manipulated within sites with throughfall reduction roofs. Seedling establishment and growth were principally affected by the spring weather in the year of emergence. In years with average to positive spring water balance, seedlings grown at the site with the highest water holding capacity had 2–5 times more aboveground biomass than seedlings grown at sites with less favourable soils. Effects of seed origin were marginal and only detectable at the drier sites: contrary to our expectations, seedlings from the Central Alpine Rhone valley, where the climatic spring water deficit is large, outperformed those from the Mediterranean. Consequently, plantation of non-native populations from dryer origin will mitigate the effects of increased summer drought at driest sites only, while the inter-annual variability of spring precipitation will continue to enable temperate conifers to regenerate on a wide range of forest soils independent of seed origin.     

缓慢干旱下春小麦叶片质膜脂肪酸组成、H~+-ATPase肥及5′-AMPase活力的变化

研究了田间缓慢干旱胁迫下,抗旱性不同的两个小麦(Triticum aestivum)品种的生长状况、质膜极性脂脂肪酸组成以及质膜关键酶活力的变化。在小麦生长发育的早期,干旱胁迫使其叶片质膜极性脂脂肪酸不饱和度下降、质膜微囊消耗O_2的速率升高、膜蛋白含量降低、H~+-ATPase(EC 3.6.1.35)活力下降、5′-AMPase(EC 3.1.3.5)活力大幅度升高;在小麦发育的后期,随着干旱的持续,小麦叶片质膜的极性脂脂肪酸不饱和度不变或升高、质膜微囊消耗O_2的速率降低、膜蛋白含量与H~+-ATPase活力升高、5′-AMPase活力下降。以上结果表明,小麦在发育的早期阶段对干旱较敏感,其细胞质膜流动性降低、细胞中能荷贮备降低;而在后期,则又表现出对干旱的适应。这些结果将有助于阐明自然干旱条件下植物的抗旱机制。     

缓慢干旱下春小麦叶片质膜脂肪酸组成、H+-ATPase及5′-AMPase活力的变化

 研究了田间缓慢干旱胁迫下,抗旱性不同的两个小麦(Triticum aestivum)品种的生长状况、质膜极性脂脂肪酸组成以及质膜关键酶活力的变化。在小麦生长发育的早期,干旱胁迫使其叶片质膜极性脂脂肪酸不饱和度下降、质膜微囊消耗O2的速率升高、膜蛋白含量降低、H+-ATPase (EC 3.6.1.35)活力下降、5'-AMPase (EC 3.1.3.5)活力大幅度升高;在小麦发育的后期,随着干旱的持续,小麦叶片质膜的极性脂脂肪酸不饱和度不变或升高、质膜微囊消耗O2的速率降低、膜蛋白含量与H+-ATPase活力升高、5'-AMPase活力下降。以上结果表明,小麦在发育的早期阶段对干旱较敏感,其细胞质膜流动性降低、细胞中能荷贮备降低;而在后期,则又表现出对干旱的适应。这些结果将有助于阐明自然干旱条件下植物的抗旱机制。