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1.
Foliar plasticity in response to ontogeny, location within the plant and environmental changes is widespread among long-lived organisms. To quantify the phenotypic variation in needle morphology and anatomy in response to a climate gradient, we compared contrasted populations of Pinus canariensis grown in five sites inside and outside the natural distribution area of the species. Most needle and growth traits were strongly affected by site. In general, site xericity increased the relative area of the dermal and transfusion tissues and decreased mesophyll and endodermis. Within each site, provenances from less productive locations tended to show longer needles, less relative area of dermal tissues but higher relative area of mesophyll and transfusion tissue than provenances from fertile origins. Although sclerophylly increased with aridity, no genetic differences were found for this trait thus apparently the ontogenetic delay of some provenances in xeric environments was not related with the formation of tougher needles. Several patterns of phenotypic response to different environments were shown. In general, all traits were plastic but the degree of plasticity was higher in traits related with growth than foliar traits. These results, combined with formerly published research, suggest that highly plastic populations rather than narrowly specialized ones have been selected in this species to cope with the complex interaction of environmental factors in its habitat.  相似文献   

2.
Pinus species show remarkable ontogenetic differences in needle morphology (heterophylly) between juvenile and adult vegetative phases. This developmental shift may play an adaptative role in their success under diverse habitats. As a first step to know the functional differences between each vegetative phase, we compared water loss through the cuticles of juvenile and adult needles of 21-month-old nursery-grown seedlings of nine hard pine species. Cuticular transpiration (CT), calculated after complete stomatal closure, was obtained by leaf-drying curves, and was related to leaf, ontogenetic and climatic parameters. The rate of cuticular transpiration (RCT) between juvenile and adult needles differed across pine species, and in particular segregated the Mediterranean species Pinus canariensis and P. halepensis, from the Eurasian P. uncinata and introduced species P. radiata. For these species, RCT was always higher in juvenile needles. The different leaf and ontogenic parameters studied were correlated with the variation in RCT among the nine pine species. We discuss this relationship in the light of the species ecology. Besides their possible adaptive interpretation, these results suggest an underlying need to consider the ontogenetic heterophylly when assessing functional traits in hard pine seedlings, in particular those traits that govern water relations.  相似文献   

3.
Effects of water stress on needle ultrastructure of 2-year-old Scots pine (Pinus sylvestris L.) and 5-year-old Norway spruce [Picea abies (L.) Karst.] seedlings were studied in greenhouse experiments. Drought stress was induced by leaving seedlings without watering, and waterlogging stress was produced by submerging the seedling containers in water. Needle samples for ultrastructural analyses were collected several times during the experiments, and samples for nutrient analyses at the end of the experiments. In drought stress, plasmolysis of mesophyll and transfusion parenchyma tissues, aggregation of chloroplast stroma and its separation from thylakoids and decreased size and abundance of starch grains in needles of both species were observed. The concentration of lipid bodies around the chloroplasts were detected in pine needles. Calcium and water concentrations in spruce needles were lower by the end of the experiments compared to controls. In waterlogging treatment, swelling of phloem cells in pine needles and large starch grains, slight swelling of thylakoids and increased translucency of plastoglobuli in chloroplasts of both species studied were observed. The phosphorus concentration in pine needles was higher while phosphorus, calcium and magnesium concentrations in spruce needles were lower in the waterlogging treatments compared to controls. Typical symptoms induced by drought stress, e. g. aggregation of chloroplast stroma and its separation from thylakoids, were detected, but, in waterlogging stress, ultrastructural symptoms appeared to be related to the developing nutrient imbalance of needles.  相似文献   

4.
Sakhalin spruce (Picea glehnii), a native species typically found in northern Japan, has been used in reforestation on hillsides exposed to strong winds. In the reforestation areas, there are south-facing (S-slope) and northwest-facing slopes (NW-slope). Climatic conditions are contrasting between the two slopes, with shallower snow cover on the S-slopes. We compared growth responses of the spruce to micro-environment between the S- and NW-slopes through soil nutrients, needle longevity, water status, photosynthesis, and nutrients in the needles. These parameters were measured in needles exposed above the snow in winter and in lower needles protected by snow cover. High-position needles suffered from drought stress, especially in winter, and needles were shed early in the year on both slopes. Low-position needles did not suffer from drought stress, and maintained a high photosynthetic rate on both slopes. However, needle longevity was reduced on the S-slope, and concentrations of nitrogen, phosphorus, and potassium in the needles decreased with needle age. Soil nutrient concentrations were low on the S-slope, which suggests that the needles on the S-slope may remobilize nutrients from aged needles to younger needles prior to shedding. This characteristic is probably an adaptation in Sakhalin spruce to poor soil conditions.  相似文献   

5.
林恬  郑怀舟  朱锦懋 《生态学报》2022,42(18):7641-7651
植物叶片中的非结构性碳水化合物(NSC)和碳(C)、氮(N)、磷(P)含量可反映植物和生态系统对水分亏缺环境的响应及适应程度。以福建省长汀县25a马尾松(Pinus massoniana)为对象,原位观测分析了3年持续100%隔离降雨对成年马尾松针叶NSC与C、N、P的影响。结果表明:(1)持续隔离降雨导致马尾松针叶NSC含量先显著增加后减少,最终导致针叶NSC含量季节变化消失;且针叶可溶性糖含量、可溶性糖/淀粉比值在后期显著增加;(2)持续隔离降雨使马尾松针叶N含量、P含量、N∶P均表现为前期与对照组差异不显著,后期显著高于对照组;(3)马尾松针叶NSC含量、可溶性糖含量、P含量、N∶P与土壤水分含量呈显著或极显著相关,且均与P含量显著或极显著相关,与N素无相关性。以上研究结果表明,随着土壤水分含量的持续减少马尾松可通过调整针叶中NSC含量的积累及分配和提高针叶N、P含量来适应缺水环境,P含量的增加对NSC含量波动及可溶性糖和淀粉的相互转化起促进作用。  相似文献   

6.
Range shifts are among the most ubiquitous ecological responses to anthropogenic climate change and have large consequences for ecosystems. Unfortunately, the ecophysiological forces that constrain range boundaries are poorly understood, making it difficult to mechanistically project range shifts. To explore the physiological mechanisms by which drought stress controls dry range boundaries in trees, we quantified elevational variation in drought tolerance and in drought avoidance‐related functional traits of a widespread gymnosperm (ponderosa pine – Pinus ponderosa) and angiosperm (trembling aspen – Populus tremuloides) tree species in the southwestern USA. Specifically, we quantified tree‐to‐tree variation in growth, water stress (predawn and midday xylem tension), drought avoidance traits (branch conductivity, leaf/needle size, tree height, leaf area‐to‐sapwood area ratio), and drought tolerance traits (xylem resistance to embolism, hydraulic safety margin, wood density) at the range margins and range center of each species. Although water stress increased and growth declined strongly at lower range margins of both species, ponderosa pine and aspen showed contrasting patterns of clinal trait variation. Trembling aspen increased its drought tolerance at its dry range edge by growing stronger but more carbon dense branch and leaf tissues, implying an increased cost of growth at its range boundary. By contrast, ponderosa pine showed little elevational variation in drought‐related traits but avoided drought stress at low elevations by limiting transpiration through stomatal closure, such that its dry range boundary is associated with limited carbon assimilation even in average climatic conditions. Thus, the same climatic factor (drought) may drive range boundaries through different physiological mechanisms – a result that has important implications for process‐based modeling approaches to tree biogeography. Further, we show that comparing intraspecific patterns of trait variation across ranges, something rarely done in a range‐limit context, helps elucidate a mechanistic understanding of range constraints.  相似文献   

7.
Needle ageing and senescence were studied in Scots pine (Pinus sylvestris) trees growing in natural conditions with minimal anthropogenic influence. The four existing needle generations were analyzed by light and transmission electron microscopy before and during autumnal yellowing of the oldest needle generation. The change from green to yellow occurred within less than 4 days in central Finland. The structure of the oldest needles remained largely intact as long as they were green. Increase in mitochondrion size and hypertrophy of the phloem parenchyma were the only changes, probably related to the approaching senescence. In the yellow needles, the structure of the mesophyll tissue varied from nearly intact with reduced chloroplasts and higher numbers of plastoglobuli, to totally disintegrated cells. In the disintegrated cells, peroxisomes were absent, and chloroplasts were smaller with a patchy appearance and degraded, eventually empty-looking stroma. Mitochondria were enlarged, but retained integrity until the last stages of deterioration, and lipids increased. At the light microscopic level, vacuolar volume in mesophyll cells and cavity formation in transfusion tissue increased. Ageing was characterized by increases in the vacuolar volume and cytoplasmic lipids, altered appearance of vacuolar tannin from homogenous ‘sandy’, to large spherical drops and finally to a large mass in the mesophyll, and by hypertrophy and tannin accumulation in the phloem parenchyma. Changes related to needle ageing, senescence and cell location in the mesophyll tissue were discussed relative to findings with stress by strong light, weather conditions and ozone.  相似文献   

8.
Tree decline and deaths have been observed among 15 to 20-year-old Scots pines (Pinus sylvestris L.) in a dry heath forest in southwestern Finland. The sudden decline in height growth, the dieback of leading shoots and the yellowing of needles in young shoots in the upper part of the tree are typical symptoms of the decline of these young pines. Needle ultrastructure and chemical composition of Scots pines with or without decline and fluctuations of them in different seasons were studied. Afflicted trees were found to suffer from a deficiency in calcium and magnesium with low concentration of foliar nitrogen and phosphorus observed in all the trees studied. Ultrastructural study revealed changes characteristic of different seasons and measured nutrient status of needles. A clear reduction of membrane system in chloroplasts, especially related to Mg deficiency, was observed in most samples. The symptoms related to N deficiency, the translucent appearance of the cytoplasm and chloroplast stroma, and the elongated chloroplasts, as well as swelling of mitochondria, indicating P deficiency, were also found in the needles sampled from this forest decline area. The present study showed that it is possible to detect specific nutrient deficiency symptoms in needle ultrastructure in field samples and for use as sensitive indicators of unbalanced nutrient status.  相似文献   

9.
Environmental conditions control physiological processes in plants and thus their growth. The predicted global warming is expected to accelerate tree growth. However, the growth response is a complex function of several processes with both direct and indirect effects. To analyse this problem we have used needle nitrogen productivity, which is an aggregate parameter for production of new foliage. Data on needle dry matter, production, and nitrogen content in needles of Scots pine ( Pinus sylvestris) and Norway spruce ( Picea abies) from a wide range of climatic conditions were collected and needle nitrogen productivities, defined as dry matter production of needles per unit of nitrogen in the needle biomass, were calculated. Our results show that the nitrogen productivity for spruce is insensitive to temperature. However, for pine, temperature affects both the magnitude of nitrogen productivity at low needle biomass and the response to self-shading but the temperature response is small at the high end of needle biomass. For practical applications it may be sufficient to use a species-specific nitrogen productivity that is independent of temperature. Because temperature affects tree growth indirectly as well as through soil processes, the effects of temperature change on tree growth and ecosystem carbon storage should mainly be derived from effects on nitrogen availability through changes in nitrogen mineralization. In addition, this paper summarises data on dry matter, production and nitrogen content of needles of conifers along a temperature gradient.  相似文献   

10.
Abstract

Observations on needle length and longevity of Pinus pinea L. - We investigated needle length and longevity of Pinus pinea L. to understand their ecological significance. Samplings regarded five coastal pinewoods (four in the province of Grosseto, one in the province of Lucca) and isolated trees in different sites of Tuscany. We found a significant variation in needle length among different positions of the crown (central, intermediate and peripheral), among individuals of the same site and among different sites; isolated trees showed longer needles and a longer retention than pines in stands. We also obtained a highly significant linear regression between length of different-aged needles (from 1988 to 1992) and rainfall of the period march-august of the same years. Our results suggest that, despite its high variability, needle ength is influenced by climatic conditions (overall by rainfalls) and by the water availability of the site where pines grow. Needle retention depends largely on the position of needles within the crown and on the drought effect: at the upper position, three or, seldom, four-year old needle can persist if the summer drought period is not too long and the site conditions are favourable; at the lower level only two-year old needles are retained. Needle longevity decreases sharply in years with severe drought, therefore it can be used as an index of climatic and edaphic stress.  相似文献   

11.
The genus Pinus has wide geographical range and includes species that are the most economically valued among forest trees worldwide. Pine needle length varies greatly among species, but the effects of needle length on anatomy, function, and coordination and trade‐offs among traits are poorly understood. We examined variation in leaf morphological, anatomical, mechanical, chemical, and physiological characteristics among five southern pine species: Pinus echinata, Pinus elliottii, Pinus palustris, Pinus taeda, and Pinus virginiana. We found that increasing needle length contributed to a trade‐off between the relative fractions of support versus photosynthetic tissue (mesophyll) across species. From the shortest (7 cm) to the longest (36 cm) needles, mechanical tissue fraction increased by 50%, whereas needle dry density decreased by 21%, revealing multiple adjustments to a greater need for mechanical support in longer needles. We also found a fourfold increase in leaf hydraulic conductance over the range of needle length across species, associated with weaker upward trends in stomatal conductance and photosynthetic capacity. Our results suggest that the leaf size strongly influences their anatomical traits, which, in turn, are reflected in leaf mechanical support and physiological capacity.  相似文献   

12.
13.
Our goal was to establish the tolerance to flooding and drought of seedlings from a hydric gradient of different seed sources to provide recommendations for forest restoration in the face of climate change. We used Drimys winteri var. chilensis, a tree species that grows from extreme arid zones to wetlands along Chile, as the study subject. We expected that seedlings of xeric origin would perform better in drought conditions than populations from moist environments, and vice versa for flooding tolerance. We collected D. winteri seeds from xeric, mesic and wet environments. Seedlings at two development stages were submitted to an extreme flooding and drought treatment during 2 or 4 months in a common garden. After the flooding and drought assays finished, the number of surviving and damaged seedlings, lenticels and adventitious root presence, height, new leaves and specific leaf area, shoot/root ratio, water potential and/or chlorophyll fluorescence (Fv/Fm), were recorded. We found that flooding and drought affected almost all the parameters studied negatively. The xeric population seedlings, at both development stages studied, were the most tolerant to the drought and, unexpectedly, also to the flooding treatment. We recommend restoring with seedlings of xeric origin especially in arid areas where sudden flooding is frequent, as occurs in the Andes Mountains. In the face of climate change, we recommend carrying out common garden and field studies before advising which population origin should be used for restoration, since they do not always respond in accordance with expected patterns of local adaptation.  相似文献   

14.
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.  相似文献   

15.
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  相似文献   

16.

Background and Aims

It is widely accepted that hydraulic failure due to xylem embolism is a key factor contributing to drought-induced mortality in trees. In the present study, an attempt is made to disentangle phenotypic plasticity from genetic variation in hydraulic traits across the entire distribution area of a tree species to detect adaptation to local environments.

Methods

A series of traits related to hydraulics (vulnerability to cavitation and hydraulic conductivity in branches), growth performance and leaf mass per area were assessed in eight Pinus canariensis populations growing in two common gardens under contrasting environments. In addition, the neutral genetic variability (FST) and the genetic differentiation of phenotypic variation (QST) were compared in order to identify the evolutionary forces acting on these traits.

Key Results

The variability for hydraulic traits was largely due to phenotypic plasticity. Nevertheless, the vulnerability to cavitation displayed a significant genetic variability (approx. 5 % of the explained variation), and a significant genetic × environment interaction (between 5 and 19 % of the explained variation). The strong correlation between vulnerability to cavitation and survival in the xeric common garden (r = –0·81; P < 0·05) suggests a role for the former in the adaptation to xeric environments. Populations from drier sites and higher temperature seasonality were less vulnerable to cavitation than those growing at mesic sites. No trade-off between xylem safety and efficiency was detected. QST of parameters of the vulnerability curve (0·365 for P50 and the slope of the vulnerability curve and 0·452 for P88) differed substantially from FST (0·091), indicating divergent selection. In contrast, genetic drift alone was found to be sufficient to explain patterns of differentiation for xylem efficiency and growth.

Conclusions

The ability of P. canariensis to inhabit a wide range of ecosystems seemed to be associated with high phenotypic plasticity and some degree of local adaptations of xylem and leaf traits. Resistance to cavitation conferred adaptive potential for this species to adapt successfully to xeric conditions.  相似文献   

17.
In many tree species, physiological adaptations to drought include the accumulation of osmotically active substances and/or the presence of particular compatible solutes, among them cyclitols. Recently, the cyclitol quercitol was identified in species of Eucalyptus, a diverse genus whose speciation is probably driven by adaptation to water availability. We subjected seedlings of 13 Eucalyptus species from different ecosystems ('mesic' and 'xeric') and different sub-generic taxonomic groups to 10 weeks of water deficit (WD) treatment. Pre-dawn water potentials (psi(pdwn)) and relative water content (RWC) were determined in shoots, and total osmolality, soluble low-molecular-weight carbohydrates and cyclitols were measured in leaves and roots. Responses to water deficit followed two distinct patterns: Eucalyptus species from 'mesic' environments adjusted concentrations of sucrose (through increased levels of sucrose and decreases in RWC) in response to water deficit, whereas 'xeric' species increased concentrations of quercitol (through reductions in RWC). In root tissues, only species from xeric environments contained high levels of quercitol and mannitol, increasing under WD conditions. We suggest that the former (mesic) strategy may be beneficial to respond to short-lasting drought conditions, because sucrose is easily metabolized, whereas the latter (xeric) strategy may relate to an effective acclimation to longer-lasting drought. These physiological response groups are also related to taxonomic groups within the genus.  相似文献   

18.
Drought is one of the critical factors limiting plant growth, and knowledge about genetic adaptation to drought is necessary to develop strategies for successful reforestation on drought-prone sites. In this study, genetic variation was investigated in 98 full- and half-sib radiata pine (Pinus radiata D. Don) families, representing Chilean coastal and interior populations, subjected to two water regimes: well-watered and drought-stressed. Assessed traits, in 5-month-old seedlings, included height (H), diameter (D), height-to-diameter ratio (HDR), dry biomass of needles (NDW), stem (SDW), roots (RDW), and total (TDW), root-to-shoot ratio (RSR), and survival (SUR). After 115 days of treatment, growth, biomass, and survival were nearly two times higher under the well-watered regime than under the stressed one. Families differed significantly in most traits, with individual tree heritabilities ranging from 0.14 for SUR to 0.63 for D in the well-watered treatment. Families from the interior showed the highest heritability for D, SDW, RDW, and TDW when grown in the water-stress treatment. The genetic correlations between treatments were moderately strong, which suggests the presence of a genotype by watering regime interaction. Most traits were strongly correlated (genetic correlations often exceeded 0.40). Compared to the first generation families from coastal sites, the third generation families from the interior sites showed an increase in SUR and RSR. Thus, potential exists to screen families at the seedling stage for drought hardiness and to identify parents from the interior sites with potential to produce a more drought-resistant breed with satisfactory growth rates and yields in dry environments.  相似文献   

19.
Reliable and objective estimations of specific leaf area (SLA) and leaf area index (LAI) are essential for accurate estimates of the canopy carbon gain of trees. The variation in SLA with needle age and position in the crown was investigated for a 73-year-old Scots pine (Pinus sylvestris L.) stand in the Belgian Campine region. Allometric equations describing the projected needle area of the entire crown were developed, and used to estimate stand needle area. SLA (cm2 g−1) as significantly influenced by the position in the crown and by needle age (current-year versus 1-year-old needles). SLA increased significantly from the top to the bottom of the crown, and was significantly higher near the interior of the crown as compared to the crown edge. SLA of current-year needles was significantly higher than that of 1-year-old needles. Allometric relationships of projected needle area with different tree characteristics showed that stem diameter at breast height (DBH), tree height and crown depth were reliable predictors of projected needle area at the tree level. The allometric relationships between DBH and projected needle area at the tree level were used to predict stand-level needle area and estimate LAI. The LAI was 1.06 (m2 m−2) for current-year needles and 0.47 for 1-year-old needles, yielding a total stand LAI of 1.53.  相似文献   

20.
High-mountain-ecosystems in the Mediterranean-type climate are exceptional because of their outstanding biodiversity but also because of their characteristic drought stress in summer. Still, plant functioning in these habitats has been largely understudied. Here, morphological, photochemical, and biochemical traits were seasonally assessed in six shrubs characterized by contrasting morphological traits, in the Teide mountain in the Canary Islands. Two adjacent populations, the first located in an open site and the second in the understorey of Pinus canariensis treeline forest, were evaluated. We aimed at disentangling (1) the role of morphological and biochemical photoprotective strategies and of their seasonal plasticity to cope with changing environmental conditions in this semiarid ecosystem, (2) how the interspecific differences in biochemical photoprotection are related to leaf morphology and phenology and (3) how living in the understory of the treeline may affect those responses. Our results indicate that both morphological and biochemical traits (particularly leaf habit, morphology and carotenoids from the β-branch) play an intricate role in photoprotection, and that a high interspecific variability exists. According to the down-regulation of photochemical activity and the upregulation of photoprotective molecules, species could be grouped into three types: (1) those more responsive to summer stress (e.g. Descurainia bourgeauana); (2) those more responsive to winter stress (e.g. Pterocephalus lasiospermus, Scrophularia glabrata and Adenocarpus viscosus); and (3) those showing rather constant behavior across seasons (e.g. Spartocytisus supranubius and Erysimum scoparium). In all the species, plants in the open site showed a marked seasonal physiological response in most of the studied parameters. Pinus canariensis canopy buffers environmental abiotic constrains. On a global change scenario, and provided further functional studies are needed, our results pinpoints heterogeneity in the sensitivity of these species against for instance late-frost or summer-heat/drought events, which could easily shift current species distribution in the coming years.  相似文献   

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