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.     

Ecophysiological responses of two Mediterranean shrubs, Erica multiflora and Globularia alypum, to experimentally drier and warmer conditions

A new approach was used to experimentally dry and warm a Mediterranean shrubland. By means of automatically sliding curtains, the drought period was extended by excluding rain over the two growing seasons (spring and autumn), and passive warming was created by avoiding infra‐red dissipation at night over the whole year. The aim of the study was to test how a future extended drought period and an increase in temperatures could affect the photosynthetic and water use strategies of two co‐occurring Mediterranean shrubs, Erica multiflora and Globularia alypum, which are common species of the dry coastal shrublands. The shoot water potential, leaf gas exchange rates and chlorophyll a fluorescence of plants was monitored seasonally during two years (1999–2001). In addition we measured the photosynthetic response curves to light and CO₂ in autumn 2001 and the foliar N concentration and leaf C and N stable isotopes in summer 1999 and 2000. Droughted plants of both shrub species showed lower shoot water potentials, transpiration rates and stomatal conductances than control plants, although there was a high seasonal variability. Drought treatment reduced significantly the overall leaf net photosynthetic rates of E. multiflora, but not of G. alypum. Droughted plants of E. multiflora also showed lower leaf net photosynthetic rates in response to light and CO₂ and lower carboxylation efficiency than controls, but there was no significant effect of drought on its overall photosystem II (PSII) photochemical efficiency. Although warming treatment did not affect the leaf net photosynthetic rates of the two species overall the study, it increased significantly the carboxylation efficiency and leaf net photosynthetic rates of G. alypum plants in response to CO₂ levels in autumn 2001. In addition, warming treatment increased the potential photochemical efficiency of PSII (F_v/F_m) of both species (but especially of G. alypum) at predawn or midday and mainly in autumn and winter. Thus, the results suggest that drier conditions might decrease the annual productivity of these Mediterranean shrubs, particularly of E. multiflora, and that future warming could alleviate the present low temperature constraints of the photosynthetic performance of the two studied species, but especially of G. alypum, during the colder seasons. Ultimately, drier and warmer conditions in the near future may change the competitive relationship among these species in such Mediterranean ecosystems.     

Drought advances spring growth phenology of the Mediterranean shrub Erica multiflora

Current climate projections predict drier and warmer conditions in the Mediterranean basin over the next century. While advanced spring growth due to warming has been described in the literature, few data are available on the effects of drought on phenology. Hence, the phenology and growth of two Mediterranean shrubs, Erica multiflora and Globularia alypum, was studied in a rainfall exclusion field experiment to simulate spring drought in a natural shrubland. We estimated the onset of growth in spring by monitoring the appearance of new stems, and the end of growth in summer by following the elongation of stems. Drought treatment caused earlier onset of the spring growing season in E. multiflora, whereas no advance was observed in G. alypum. However, growth cessation was not affected in E. multiflora. Drought reduced the growth of both shrubs, as reflected in less stem elongation. The results show that a drier climate might affect not only growth but also spring phenology of some Mediterranean species. We suggest that a reduction in the cooling effect of transpiration may have analogous effects to warming and might advance the start of growth in E. multiflora, a species whose phenology has been described as warming‐sensitive. The lengthening of the growing season resulting from advanced growth did not imply higher productivity, as growth was restricted by drought.     

Partitioning of water and nitrogen in co-occurring Mediterranean woody shrub species of different evolutionary history

We studied the interspecific and intraspecific variation in the development of water stress and in the use of different water and nitrogen sources during the spring (wet season) and summer (dry season) in a shrub community in NE Spain. We measured shoot water potentials, stable deuterium isotopic composition (D) of xylem sap, leaf mass per area, leaf N and C concentrations, gas exchange, leaf ¹³C, and leaf ¹⁵N of the dominant species (Quercus coccifera, Arbutus unedo, Pistacia lentiscus, Erica multiflora, Globularia alypum). The D, the ¹³C and the shoot water potential values showed diurnal, seasonal, intraspecific and interspecific variation in the source and use of water. There was also seasonal, intraspecific and interspecific variation in the foliar ¹⁵N and N concentrations. In summer, some species (A. unedo, P. lentiscus and E. multiflora) presented significantly different D values in morning and afternoon measurements likely indicating that they used different sources of water during the day, and a dual root system in these species. We conjecture that dew may be one of these water sources. Species predawn water potential was negatively correlated with species xylem water D. There was also a positive correlation between ¹³C and D in P. lentiscus, species for which we took additional samples from nearby sites. These results suggest that the access to water from greater depths allowed the maintenance of more favourable plant water supply. Multivariate principal component analysis based on the studied hydrological and isotope variables clearly separated the seasons (wet spring and dry summer) and the species. The species resulted separated according to their evolutionary history (Pre-Mediterranean and Mediterranean) and the associated root and functional traits. These results show water (and nitrogen) partitioning among coexisting species of the same functional type (Mediterranean woody shrubs). They also show the great intraspecific plasticity of responses to resource availability.     

Gas exchange and water relations of three <Emphasis Type="Italic">Vitis vinifera</Emphasis> L. cultivars growing under Mediterranean climate

Optical characteristics, contents of photosynthetic pigments, total soluble sugars, and starch, rates of gas exchange, chlorophyll (Chl) a fluorescence, and leaf water relations were analysed in three Vitis vinifera L. cultivars, Tinto Cão (TC), Touriga Nacional (TN), and Tinta Roriz (TR), grown in Mediterranean climate. Chl content was significantly lower in TC than in TN and TR leaves, while the Chl a/b ratio was higher. TR had the lowest net photosynthetic rate, stomatal conductance, and contents of soluble sugars and starch than TN and TC. In spite of low Chl content, TC showed the lowest photon absorbance and the highest photochemical efficiency of photosystem 2. TC had the lowest predawn and midday leaf water potential. The capability for osmotic adjustment was similar among cultivars and the calculated modulus of elasticity was higher in TC leaves. The typical lighter green leaves of TC seemed to be an adaptive strategy to high irradiance and air temperature associated to water stress.     

Leaf shape linked to photosynthetic rates and temperature optima in South African <Emphasis Type="Italic">Pelargonium</Emphasis> species

The thermal response of gas exchange varies among plant species and with growth conditions. Plants from hot dry climates generally reach maximal photosynthetic rates at higher temperatures than species from temperate climates. Likewise, species in these environments are predicted to have small leaves with more-dissected shapes. We compared eight species of Pelargonium (Geraniaceae) selected as phylogenetically independent contrasts on leaf shape to determine whether: (1) the species showed plasticity in thermal response of gas exchange when grown under different water and temperature regimes, (2) there were differences among more- and less-dissected leafed species in trait means or plasticity, and (3) whether climatic variables were correlated with the responses. We found that a higher growth temperature led to higher optimal photosynthetic temperatures, at a cost to photosynthetic capacity. Optimal temperatures for photosynthesis were greater than the highest growth temperature regime. Stomatal conductance responded to growth water regime but not growth temperature, whereas transpiration increased and water use efficiency (WUE) decreased at the higher growth temperature. Strikingly, species with more-dissected leaves had higher rates of carbon gain and water loss for a given growth condition than those with less-dissected leaves. Species from lower latitudes and lower rainfall tended to have higher photosynthetic maxima and conductance, but leaf dissection did not correlate with climatic variables. Our results suggest that the combination of dissected leaves, higher photosynthetic rates, and relatively low WUE may have evolved as a strategy to optimize water delivery and carbon gain during short-lived periods of high soil moisture. Higher thermal optima, in conjunction with leaf dissection, may reflect selection pressure to protect photosynthetic machinery against excessive leaf temperatures when stomata close in response to water stress.     

Water-use strategies of six co-existing Mediterranean woody species during a summer drought

Drought stress is known to limit plant performance in Mediterranean-type ecosystems. We have investigated the dynamics of the hydraulics, gas exchange and morphology of six co-existing Mediterranean woody species growing under natural field conditions during a drought that continued during the entire summer. Based on the observed minimum leaf water potentials, our results suggest that the six co-existing species cover a range of plant hydraulic strategies, from isohydric to anisohydric. These differences are remarkable since the selected individuals grow within several meters of each other, sharing the same environment. Surprisingly, whatever the leaf water potentials were at the end of the dry period, stomatal conductance, photosynthesis and transpiration rates were relatively similar and low across species. This result contradicts the classic view that anisohydric species are able to maintain gas exchange for longer periods of time during drought stress. None of the plants showed the expected structural acclimation response to the increasing drought (reduction of leaf-to-sapwood area ratio), thereby rejecting the functional equilibrium hypothesis for our study system. Instead, three of the six species increased photosynthetic area at the branch level. The observed dissimilar patterns of gas exchange, hydraulics and morphology across species seem to be equally successful given that photosynthesis at the leaf level was maintained at similar rates over the whole dry period.     

Physiological responses of beech and sessile oak in a natural mixed stand during a dry summer

Responses of CO2 assimilation and stomatal conductance to decreasing leaf water potential, and to environmental factors, were analysed in a mixed natural stand of sessile oak (Quercus petraea ssp. medwediewii) and beech (Fagus svlvatica L.) in Greece during the exceptionally dry summer of 1998. Seasonal courses of leaf water potential were similar for both species, whereas mean net photosynthesis and stomatal conductance were always higher in sessile oak than in beech. The relationship between net photosynthesis and stomatal conductance was strong for both species. Sessile oak had high rates of photosynthesis even under very low leaf water potentials and high air temperatures, whereas the photosynthetic rate of beech decreased at low water potentials. Diurnal patterns were similar in both species but sessile oak had higher rates of CO2 assimilation than beech. Our results indicate that sessile oak is more tolerant of drought than beech, due, in part, to its maintenance of photosynthesis at low water potential.     

A preliminary study of the responsiveness to seasonal atmospheric and rainfall patterns of wash woodland species in the arid Richtersveld

Seasonal carbon and water relations were compared among seven tree or shrub wash woodland species in the winter rainfall desert of the Richtersveld National Park, South Africa. Plants were generally aseasonal with respect to gas exchange, but responsive to rainfall events with respect to water relations and phenology. Relatively narrow annual ranges in potential evapotranspiration due to the maritime influence could explain why these plants respond more to fluctuations in water acquisition potential than to evaporative demand. Two species were summer-deciduous, but one of them (Ozoroa concolor) responded to aseasonal summer rainfall by leafing out and flowering. These two species had high shoot xylem water potentials when in leaf. All other species were sclerophyllous evergreens with low water potentials, particularly the shallow-rooted shrub Zygophyllum prismatocarpum, and Boscia albitrunca which may have a different rooting pattern to the other phreatophytes. The latter species was also unique due to its high leaf nitrogen contents, photosynthetic rates and stomatal conductances, despite very low leaf water potentials. Leaf stable carbon isotope composition C¹³C) varied between species (–22 to –27), but was lower than the mean for arid regions worldwide. The values indicated moderately high levels of water use efficiency, but a less conservative strategy in two species, including Boscia albitrunca. The affinities of these species to summer rainfall biomes, their apparent decline in the western arid regions in recent geological history following aridification, and their absence southwards in the winter rainfall regions, suggest that these wash species rely on sporadic summer rainfall events to some extent. They may be at risk if predicted increases in temperature and changes in rainfall patterns alter their effective moisture availability.     

How do Mediterranean shrub species cope with shade? Ecophysiological response to different light intensities

• Under natural conditions, light exposure for Mediterranean shrubs can be highly variable, especially during cloudy days or under a canopy, and can interfere with other environmental factors such as temperature and water availability.
• With the aim of decoupling the effect of radiation and temperature from water availability, we conducted an experiment where two perennial and three summer semi‐deciduous shrub species were subjected to different levels of irradiation. In order to follow plant responses to light exposure, we measured gas exchange, photosystem II photochemical efficiency, photosynthetic pigments and leaf mass area in spring and summer.
• Results showed that all study species presented a plastic response to different light conditions, and that light‐related traits varied in a coordinated manner. Summer semi‐deciduous species exhibited a more opportunistic response, with higher photosynthesis rates in full sun, but under shade conditions, the two strategies presented similar assimilation rates. Stomatal conductance did not show such a drastic response as photosynthetsis, being related to changes in WUE. Daily cycles of F_v/F_m revealed a slight photoinhibitory response during summer, mainly in perennial species. In all cases photosynthetic pigments adjusted to the radiation level; leaves had lower chlorophyll content, higher pool of xanthophylls and higher proportion of the de‐epoxydaded state of xanthophylls under sun conditions. Lutein content increased in relation to the xanthophyll pool under shade conditions.
• Our results evidenced that radiation is an important driving factor controlling morphological and physiological status of Mediterranean shrub species, independently of water availability. Summer semi‐deciduous species exhibit a set of traits with higher response variability, maximising their photosynthetic assimilation under different sun conditions.

     

Differential uptake of summer precipitation among co-occurring trees and shrubs in a pinyon-juniper woodland

Measurements of the ratio of deuterium to hydrogen (D/H) in stem xylem water were used to determine the relative uptake of summer precipitation by four co-occurring plant species in southern Utah. The species compared included two trees, Juniperus osteosperma and Pinus edulis, and two shrubs, Artemisia tridentata and Chrysothamnus nauseousus. There were significant differences among species in the relative use of summer precipitation. Chrysothamnus nauseosus had stem water D/H ratios in May through August 1990 that were not significantly different from that of groundwater. In contrast, the other three species had stem water D/H ratios that were intermediate between the groundwater value and summer precipitation values, indicating that a mixture of both precipitation and groundwater was being used by these species. The two tree species generally had higher D/H values than did A. tridentata indicating a higher average uptake of summer precipitation, although the roots of J. osteosperma and P. edulis may not be as responsive to small precipitation events as A. tridentata. There was a strong negative correlation between stem water D/H ratios and predawn water potential, which suggests a relationship between plant rooting pattern and water source use. In addition, water-use efficiency during photosynthetic gas exchange, calculated from leaf carbon isotope composition, differed among species and was strongly correlated with differences in the relative uptake of summer precipitation.     

Daily sap flow rate as an indicator of drought avoidance mechanisms in five Mediterranean perennial species in semi-arid southeastern Spain

Daily sap flow rate was determined in five Mediterranean species (Pinus halepensis, Quercus coccifera, Pistacia lentiscus, Erica multiflora, and Stipa tenacissima) under two slope aspects (north- and south-facing) in a semi-arid area (Alicante, SE Spain). Sap flow velocity was measured in January, May, August and October of two consecutive years (1998 and 1999) using the stem heat balance (SHB) method. Our results have demonstrated the effects of global radiation (R _g), vapour pressure deficit (VPD) on the sap flow velocity per unit of leaf area. Mean daily sap flow rates (Q _md) showed values between 0.001 and 0.202 g H₂O cm⁻² leaf area day⁻¹. Q _md values were higher on the south-facing slope than on the north-facing slope. In most species, the Q _md was higher in 1998 than in 1999 due to the higher soil water content, temperature and VPD in 1998. In all five species, a decrease in predawn leaf water potential was accompanied by a decrease in mean daily sap flow rates; nevertheless, the responses of the five species to water deficit conditions were different. In this context, we have linked the drought avoidance mechanisms of the different species through the combined use of daily sap flow rate and predawn leaf water potential under different water deficit conditions. We conclude that Pinus halepensis, Pistacia lentiscus and Erica multiflora show water-savers mechanisms to cope with drought, while Quercus coccifera and Stipa tenacissima show water-spenders mechanisms.     

Tropic leaf movements, photosynthetic gas exchange, leaf δ13C and chlorophyll a fluorescence of three soybean species in response to water availability

Midday leaf angle, photosynthetic gas exchange, stable carbon isotope ratio ( δ ¹³C), and chlorophyll a fluorescence among three wild soybeans, Glycine soja, G . tomentella and G . tabacina, from habitats with different water availability were examined. Plants grown under low water availability had reduced leaf area, photosynthetic and electron transport rates, more positive δ ¹³C values, and more vertical midday leaf angles. The three species differed in midday leaflet orientation, leaf size, photosynthesis and fluorescence responses to water availability. The species differences were consistent with the water availability of their habitat. G . soja , which grows in the wettest habitats, was shown to be the most photosynthetically susceptible to low water treatment and tended to have the most vertical midday leaflet angles. In contrast, G . tabacina , distributed in the driest habitats, had the least vertical midday leaflet angles and the lowest photosynthetic sensitivity to low water availability. G . tomentella , inhabiting an intermediate habitat, had intermediate midday leaf angles and photosynthetic responses. Our results support the hypothesis that paraheliotropic leaf movements respond in concert with photosynthetic characteristics in soybean leaves such that water use efficiency is enhanced and the risk of photoinhibition under water stress conditions is reduced.     

Effects of experimental warming and drought on biomass accumulation in a Mediterranean shrubland

We studied the effects of experimental warming and drought on the plant biomass of a Mediterranean shrubland. We monitored growth at plant level and biomass accumulation at stand level. The experimentation period stretched over 7 years (1999–2005) and we focused on the two dominant shrub species, Erica multiflora L. and Globularia alypum L. and the tree species Pinus halepensis L. The warming treatment increased shoot elongation in E. multiflora, and the drought treatment reduced shoot elongation in G. alypum. The elongation of P. halepensis remained unaffected under both treatments. The balance between the patterns observed in biomass accumulation for the three studied species in the drought plots (reduction in E. multiflora and P. halepensis and increase in G. alypum) resulted in a trend to reduce 33% the biomass of the drought treatment plots with respect to the untreated plots, which almost doubled their biomass from 1998 to 2005. The results also suggest that under drier conditions larger accumulation of dead biomass may occur at stand level, which combined with higher temperatures, may thus increase fire risk in the Mediterranean area.     

Regulation and acclimation of leaf gas exchange in a piñon–juniper woodland exposed to three different precipitation regimes

Leaf gas‐exchange regulation plays a central role in the ability of trees to survive drought, but forecasting the future response of gas exchange to prolonged drought is hampered by our lack of knowledge regarding potential acclimation. To investigate whether leaf gas‐exchange rates and sensitivity to drought acclimate to precipitation regimes, we measured the seasonal variations of leaf gas exchange in a mature piñon–juniper Pinus edulis–Juniperus monosperma woodland after 3 years of precipitation manipulation. We compared trees receiving ambient precipitation with those in an irrigated treatment (+30% of ambient precipitation) and a partial rainfall exclusion (?45%). Treatments significantly affected leaf water potential, stomatal conductance and photosynthesis for both isohydric piñon and anisohydric juniper. Leaf gas exchange acclimated to the precipitation regimes in both species. Maximum gas‐exchange rates under well‐watered conditions, leaf‐specific hydraulic conductance and leaf water potential at zero photosynthetic assimilation all decreased with decreasing precipitation. Despite their distinct drought resistance and stomatal regulation strategies, both species experienced hydraulic limitation on leaf gas exchange when precipitation decreased, leading to an intraspecific trade‐off between maximum photosynthetic assimilation and resistance of photosynthesis to drought. This response will be most detrimental to the carbon balance of piñon under predicted increases in aridity in the southwestern USA.     

Grasstree (Xanthorrhoea preissii ) leaf growth in relation to season and water availability

Abstract Water stress usually arrests growth of even the most deep‐rooted species during summer drought in Mediterranean‐type climates. However, scant evidence suggests that grasstrees may represent an unusual exception. We used weather data and plant water potential to investigate the relationship between leaf growth and season in the grasstree, Xanthorrhoea preissii Endl. (Xanthorrhoeaceae). Leaf production in two contrasting habitats revealed continuous annual growth, oscillating between maximum rates (2.5–3.2 leaves/d) in late‐spring to autumn, to a minimum rate of 0.5 leaf/d during winter but never stopping. While the rate of leaf production during the fast‐growth season was positively correlated with temperature above 17–18°C, leaf elongation commenced substantially earlier in the year (from 12°C). Leaf water potentials cycled annually, with predawn readings commonly measured as zero during winter–spring and as low as ?1.26 MPa during summer, but never indicating stress by exceeding the turgor loss point. Leaf death was synchronized with summer drought. The fast (summer) growth period was characterized by rapidly fluctuating leaf production, particularly in banksia woodland, where plant growth reliably responded quickly to >18 mm of rainfall. Within 24 h of 59 mm of simulated rainfall, grasstrees in banksia woodland showed a marked increase in water potential, and leaf production reached 7.5 times the controls, confirming their capacity to respond to temporary spasmodic summer rains. Rainfall was the best climatic variable for predicting woodland grasstree leaf production during summer, whereas leaf production of forest grasstrees was most closely correlated with daylength. This plastic response of grasstrees between seasonal weather extremes is relatively rare among other mediterranean floras, and has implications for a recently proposed technique for ageing grasstrees.     

Drought controls over conductance and assimilation of a Mediterranean evergreen ecosystem: scaling from leaf to canopy

Drought control over conductance and assimilation was assessed using eddy flux and meteorological data monitored during four summer periods from 1998 to 2001 above a closed canopy of the Mediterranean evergreen oak tree Quercus ilex. Additional discrete measurements of soil water content and predawn leaf water potential were used to characterize the severity of the drought. Canopy conductance was estimated through the big‐leaf approach of Penman–Monteith by inverting latent heat fluxes. The gross primary production ( GPP ) was estimated by adding ecosystem respiration to net ecosystem exchange. Ecosystem respiration was deduced from night flux when friction velocity ( u _*) was greater than 0.35 m s^?1. Empirical equations were identified that related maximal canopy conductance and daily ecosystem GPP to relative soil water content ( RWC) , the ratio of current soil water content to the field capacity, and to the predawn leaf water potential. Both variables showed a strong decline with soil RWC for values lower than 0.7. The sharpest decline was observed for GPP . The curves reached zero for RWC =0.41 and 0.45 for conductance and GPP , respectively. When the predawn leaf water potential was used as a surrogate for soil water potential, both variables showed a hyperbolic decline with decreasing water potential. These results were compared with already published literature values obtained at leaf level from the same tree species. Scaling up from the leaf to ecosystem highlighted the limitation of two big‐leaf representations: Penman–Monteith and Sellers' Π factor. Neither held completely for comparing leaf and canopy fluxes. Tower measurements integrate fluxes from foliage elements clumped at several levels of organization: branch, tree, and ecosystem. The Q. ilex canopy exhibited non‐random distribution of foliage, emphasizing the need to take into account a clumping index, the factor necessary to apply the Lambert–Beer law to natural forests. Our results showed that drought is an important determinant in water losses and CO₂ fluxes in water‐limited ecosystems. In spite of the limitations inherent to the big‐leaf representation of the canopy, the equations are useful for predicting the influence of environmental factors in Mediterranean woodlands and for interpreting ecosystem exchange measurements.     

四种不同生活型植物幼苗对喀斯特生境干旱的生理生态适应性

喀斯特石漠化是我国西南喀斯特地区最严重的生态环境问题, 生境干旱是限制该地区植物生长的主要因素之一, 掌握喀斯特植被不同演替阶段不同生活型植物对干旱胁迫的适应策略有助于提高植被恢复的成功率。通过人工模拟4种干旱强度, 测定叶片水势、气体交换、叶绿素荧光、光合色素含量、渗透调节物质浓度、抗氧化酶活性以及生物量, 研究了喀斯特地区4种不同生活型植物幼苗对干旱胁迫的适应策略。这4种植物为常绿灌木火棘(Pyracantha fortuneana)、落叶灌木小果蔷薇(Rosa cymosa)、常绿乔木猴樟(Cinnamomum bodinieri)和落叶乔木圆果化香树(Platycarya longipes)。结果表明: 随着干旱程度的加深, 4种植物幼苗的叶片水势、光合能力、叶绿素含量、生物量增长、叶重比(LMR)、叶面积比(LAR)和比叶面积(SLA)逐渐下降, 而热耗散(NPQ)、类胡萝卜素与叶绿素含量比值、丙二醛含量和根重比(RMR)逐渐上升; 圆果化香树和猴樟的水分利用效率(A_n/g_s)、渗透调节物质浓度和抗氧化酶活性呈先升高后降低的趋势, 而火棘和小果蔷薇的A_n/g_s、脯氨酸含量和超氧化物歧化酶活性呈上升趋势。严重干旱下, 火棘和小果蔷薇幼苗的叶片水势和叶绿素含量下降较少, 具有较高的光合能力和生物量增长, 这主要是由于它们具有较低的SLA和LAR、较高的NPQ和A_n/g_s以及较高的渗透调节能力和抗氧化保护能力。中度干旱下, 猴樟幼苗叶片水势下降很少, LMR和LAR也较高, 脯氨酸含量和抗氧化酶活性非常高。但在严重干旱下, 其叶片水势、LMR、LAR和生物量增长大幅度下降, 最大光化学效率和光合速率也非常低, 渗透调节能力与抗氧化酶活性大幅度下降至正常水平以下。水分好的条件下, 圆果化香树幼苗具有较高的RMR以吸收充足的水分, 具有较高的LAR和叶绿素含量, 保证了生物量的大量积累。然而, 干旱胁迫致使其生物量大幅度下降, 主要是由于LMR、LAR、气体交换和叶绿素含量的大量下降以减少蒸腾面积、水分散失和对光能的吸收。研究结果表明, 火棘、小果蔷薇和猴樟幼苗主要采用耐旱策略, 其中猴樟抗严重干旱的能力较弱; 圆果化香树幼苗对干旱胁迫更为敏感, 主要采取避旱策略。     

Hydraulic properties and photosynthetic rates in co-occurring lianas and trees in a seasonal tropical rainforest in southwestern China

In this study, we examined wood anatomy, hydraulic properties, photosynthetic rate, and water status and osmotic regulation in three liana species and three tree species co-occurring in a seasonal tropical rain forest. Our results showed that the three liana species had larger vessel diameter, lower sapwood density, and consequently higher branch sapwood specific hydraulic conductivity (K _S) than the three tree species. Across species, K _S was positively correlated with leaf nitrogen concentration and maximum net CO₂ assimilation rate. However, it was also positively correlated with xylem water potential at 50% loss of hydraulic conductivity, indicating a trade-off between hydraulic efficiency and safety. Compared to the tree species, the liana species had higher predawn leaf water potential and lower osmotic adjustment in the dry season. The combination of more efficient water transport, higher photosynthetic rates, and their ability to access to more reliable water source at deeper soil layers in the dry season in the lianas should contribute to their fast growth.     

Leaf Gas Exchange in Canopy Species of a Venezuelan Cloud Forest

Tropical cloud forests are considered humid ecosystems with frequent cloud cover down to the ground surface. However, seasonal variation in precipitation may induce short-term water stress. For canopy leaves, this water stress may also be a consequence of large atmospheric vapor pressure deficits. The objective of this work was to study five canopy cloud forest species to determine if there are restrictions to leaf gas exchange as a consequence of seasonality in precipitation and to daily water deficit due to air evaporative demand mainly during maximum incoming radiation hours. Seasonal daily courses of microclimatic variables (air temperature, relative humidity, photosynthetic photon flux density) and plant responses (leaf water potential, stomatal conductance, CO₂ assimilation rates, leaf nitrogen concentration) were measured at 2400 m asl in Monterrey, an intermontane valley of the Venezuelan Andes. A gradient in terms of responses to water stress conditions was observed between the species, with Clusia multiflora (a 46% reduction in stomatal conductance between seasons) as the most affected and Miconia resimoides (increased stomatal conductance) responding more favorably to slight water stress conditions. If we consider the limitations of water stress and/or light conditions on CO₂ assimilation we may arrange the species into those in which water stress conditions have a greater impact on leaf carbon gain, those where light conditions are determinant and one in which both water stress and light conditions may affect leaf carbon assimilation.