Impact of extreme pre-monsoon drought on xylogenesis and intra-annual radial increments of two tree species in a tropical montane evergreen broad-leaved forest, southwest China.

Tropical montane evergreen broad-leaved forests cover the majority of forest areas and have high carbon storage in Xishuangbanna, southwest China. However, stem radial growth dynamics and their correlations with climate factors have never been analyzed in this forest type. By combining bi-weekly microcoring and high-resolution dendrometer measurements, we monitored xylogenesis and stem radius variations of the deciduous species Betula alnoides and the evergreen species Schima wallichii. We analyzed the relationships between weekly climate variables prior to sampling and the enlarging zone width or wall thickening zone width, as well as weekly radial increments and climate factors during two consecutive years (2020-2021) showing contrasting hydrothermal conditions in the pre-monsoon season. In the year 2020, which was characterized by a warmer and drier pre-monsoon season, the onset of xylogenesis and radial increments of B. alnoides and S. wallichi were delayed by three months and one month, respectively, compared to the year 2021. In 2020, xylem formation and radial increments were significantly reduced for B. alnoides, but not for S. wallichill. The thickness of enlarging zone and wall thickening zone in S. wallichill were positively correlated with relative humidity, minimum and mean air temperature, but were negatively correlated with vapor pressure deficit during 2020-2021. The radial increments of both species showed significant positive correlations with precipitation and relative humidity, and negative correlations with vapor pressure deficit and maximum air temperature during two years. Our findings reveal that drier pre-monsoon conditions strongly delay growth initiation and reduce stem radial growth, providing deep insights to understand tree growth and carbon sequestration potential in tropical forests under a predicted increase in frequent drought events.

Effect of life cycle and venation pattern on the coordination between stomatal and vein densities of herbs.

Life cycle (annual vs perennial) and leaf venation pattern (parallel and reticular) are known to be related to water use strategies in herb species and critical adaptation to certain climatic conditions. However, the effect of these two traits and how they influence the coordination between vein density (vein length per area, VLA) and stomatal density (SD) remains unclear. In this study, we examined the leaves of 53 herb species from a subtropical botanical garden in Guangdong Province, China, including herbs with different life cycles and leaf venation patterns. We assessed 21 leaf water-related functional traits for all species, including leaf area (LA), major and minor VLA, major and minor vein diameter (VD), SD and stomatal length (SL). The results showed no significant differences in mean SD and SL between either functional group (parallel venation vs reticular venation and annual vs perennial). However, parallel vein herbs and perennial herbs displayed a significantly higher mean LA and minor VD, and lower minor VLA compared to reticular vein herbs and annual herbs, respectively. There was a linear correlation between total VLA and SD in perennial and reticular vein herbs, but this kind of correlation was not found in annual and parallel vein herbs. The major VLA and minor VD were significantly affected by the interaction between life cycle and leaf venation pattern. Our findings suggested that VLA, rather than SD, may serve as a more adaptable structure regulated by herbaceous plants to support the coordination between leaf water supply and demand in the context of different life cycles and leaf venation patterns. The results of the present study provide mechanistic understandings of functional advantages of different leaf types, which may involve in species fitness in community assembly and divergent responses to climate changes.

Effect of leaf phenology and morphology on the coordination between stomatal and minor vein densities.

Leaf phenology (evergreen vs. deciduous) and morphology (simple vs. compound) are known to be related to water use strategies in tree species and critical adaptation to certain climatic conditions. However, the effect of these two traits and their interactions on the coordination between minor vein density (MVD) and stomatal density (SD) remains unclear. In this study, we examined the leaves of 108 tree species from plots in a primary subtropical forest in southern China, including tree species with different leaf morphologies and phenologies. We assessed nine leaf water-related functional traits for all species, including MVD, SD, leaf area (LA), minor vein thickness (MVT), and stomatal length (SL). The results showed no significant differences in mean LA and SD between either functional group (simple vs. compound and evergreen vs. deciduous). However, deciduous trees displayed a significantly higher mean MVD compared to evergreen trees. Similarly, compound-leaved trees have a higher (marginally significant) MVD than simple-leaved trees. Furthermore, we found that leaf morphology and phenology have significantly interactive effects on SL, and the compound-leafed deciduous trees exhibited the largest average SL among the four groups. There were significant correlations between the MVD and SD in all different tree groups; however, the slopes and interceptions differed within both morphology and phenology. Our results indicate that MVD, rather than SD, may be the more flexible structure for supporting the coordination between leaf water supply and demand in different leaf morphologies and phenologies. The results of the present study provide mechanistic understandings of the functional advantages of different leaf types, which may involve species fitness in community assembly and divergent responses to climate changes.

High vapour pressure deficit enhances turgor limitation of stem growth in an Asian tropical rainforest tree.

Tropical forests are experiencing increases in vapour pressure deficit (D), with possible negative impacts on tree growth. Tree-growth reduction due to rising D is commonly attributed to carbon limitation, thus overlooking the potentially important mechanism of D-induced impairment of wood formation due to an increase in turgor limitation. Here we calibrate a mechanistic tree-growth model to simulate turgor limitation of radial stem growth in mature Toona cilitata trees in an Asian tropical forest. Hourly sap flow and dendrometer measurements were collected to simulate turgor-driven growth during the growing season. Simulated seasonal patterns of radial stem growth matched well with growth observations. Growth mainly occurred at night and its pre-dawn build-up appeared to be limited under higher D. Across seasons, the night-time turgor pressure required for growth was negatively related to previous midday D, possibly due to a relatively high canopy conductance at high D, relative to stem rehydration. These findings provide the first evidence that tropical trees grow at night and that turgor pressure limits tree growth. We suggest including turgor limitation of tree stem growth in models also for tropical forest carbon dynamics, in particular, if these models simulate effects of warming and increased frequency of droughts.

[Intra-annual radial growth of Abies georgei and Larix potaninii and its responses to environmental factors in the Baima Snow Mountain, Northwest Yunnan, China.] / æ»‡è¥¿åŒ—ç™½é©¬é›ªå±±é•¿è‹žå†·æ‰å’Œå¤§æžœçº¢æ‰å¹´å† å¾„å‘ç”Ÿé•¿åŠ¨æ€åŠå ¶å¯¹çŽ¯å¢ƒå› å­çš„å“åº”.

Using high-resolution dendrometers, we monitored the intra-annual stem radial variations of Abies georgei and Larix potaninii in the subalpine coniferous forest in Baima Snow Mountain, Northwest Yunnan Province. The seasonal dynamics of stem radial growth of both species and their responses to environmental factors were analyzed. The results showed that the stem radial growth of A. georgei and L. potaninii mainly occurred during April to August, with the maximum growth rate in June. Compared with A. georgei, L. potaninii showed an earlier start but later cessation of stem radial growth, resulting in longer growth duration. Annual radial growth and maximum radial growth rates of L. potaninii were slightly higher than those of A. georgei. Daily growth rate of A. georgei was positively correlated with precipitation, but negatively correlated with vapor pressure deficit and air temperature. Daily growth rate of L. potaninii was positively correlated with precipitation, but negatively correlated with soil volume water content and vapor pressure deficit. Radial growth of A. georgei and L. potaninii was limited by water availability, with L. potaninii being more sensitive to moisture. Under the background of global warming, the increase of plant transpiration and soil evaporation might further aggravate soil water loss and reduce water availability for plants, which would make A. georgei and L. potaninii more vulnerable to drought stress.

Leaf gas exchange and water relations of the woody desiccation-tolerant Paraboea rufescens during dehydration and rehydration.

Desiccation-tolerant (DT) plants can withstand dehydration to less than 0.1 g H2O g-1 dry weight. The mechanism for whole-plant recovery from severe dehydration is still not clear, especially for woody DT plants. In the present study, we evaluated the desiccation tolerance and mechanism of recovery for a potentially new woody resurrection plant Paraboea rufescens (Gesneriaceae). We monitored the leaf water status, leaf gas exchange, chlorophyll fluorescence and root pressure of potted P. rufescens during dehydration and rehydration, and we investigated the water content and chlorophyll fluorescence of P. rufescens leaves in the field during the dry season. After re-watering from a severely dehydrated state, leaf maximum quantum yield of photosystem II of P. rufescens quickly recovered to well-watered levels. Leaf water status and leaf hydraulic conductance quickly recovered to well-watered levels after re-watering, while leaf gas exchange traits also trended to recovery, but at a slower rate. The maximum root pressure in rehydrated P. rufescens was more than twice in well-watered plants. Our study identified P. rufescens as a new DT woody plant. The whole-plant recovery of P. rufescens from extreme dehydration is potentially associated with an increase of root pressure after rehydration. These findings provide insights into the mechanisms of recovery of DT plants from dehydration.

[Radial growth responses of four coniferous species to climate change in the Potatso National Park, China]. / æ™®è¾¾æŽªå›½å®¶å ¬å›­å››ç§é’ˆå¶æ ‘å¾„å‘ç”Ÿé•¿å¯¹æ°”å€™å› å­çš„å“åº”.

Using the principles and methods of dendrochronology, we measured tree-ring width of four dominant coniferous species, i.e., Larix potaninii var. macrocarpa, Picea brachytyla, Pinus densata, and Abies georgei, in the Potatso National Park, and established the tree-ring width resi-dual chronologies. We analyzed the correlation of tree-ring width residual chronologies with daily and monthly climate data from the Shangrila meteorological station to analyze the response of radial growth to climate factors. The results showed that L. potaninii var. macrocarpa had the highest annual growth rate, and A. georgei had the lowest. Radial growth showed species-specific responses to climate changes, with the highest sensitivity of L. potaninii var. macrocarpa and the lowest sensitivity of P. brachytyla. Ring-width chronology of A. georgei correlated positively with mean temperature during previous winter (November and December) and current summer (July). Ring-width chronology of L. potaninii var. macrocarpa correlated positively with temperature during the early-growing season (June), but negatively with precipitation and relative humidity. Ring-width chronology of P. densata correlated positively with precipitation and humidity but negatively with maximum temperature during the early-growing season (May), indicating that its radial growth was primarily influenced by water availability during the early-growing season.

[Intra-annual radial growth of Garuga floribunda in tropical seasonal rain forest and its response to environmental factors in Xishuangbanna, Southwest China]. / çƒ­å¸¦å­£èŠ‚é›¨æž—å¤šèŠ±ç™½å¤´æ ‘å¹´å† å¾„å‘ç”Ÿé•¿åŠ¨æ€åŠå ¶å¯¹çŽ¯å¢ƒå› å­çš„å“åº”.

We examined the seasonal growth dynamics of a deciduous tree species Garuga floribunda in the tropical seasonal rain forest in Xishuangbanna and monitored the stem radial growth with both high resolution dendrometer and microcoring methods. Combining with the monitoring of non-structural carbohydrates (NSCs) in stem and environmental factors, we analyzed the eco-physiological drivers underlying the seasonal cambial activity and radial growth dynamics. The results of high reso-lution dendrometer method showed that the growth of G. floribunda began at the end of May (day of year, DOY: 149.3±7.2) and ended at the end of August (DOY: 241.0±14.7) in 2020, the annual total radial growth was 3.12 mm, and the maximum growth rate was 0.04 mm·d-1. Based on the microcoring methods, we found that xylem cell enlarging started from March 9th (69.2±6.2) and cell wall thickening ended on September 19th (DOY: 262.8±2.8). The cumulative xylem radial growth was 1.76 mm, and the maximum growth rate was 0.009 mm·d-1. The daily radial growth rate of G. floribunda was significantly and positively correlated with precipitation, relative humidity, daily minimum air temperature, soil moisture and temperature at the depth of 20 cm, and was negatively correlated with daily maximum air temperature, vapor pressure deficit, maximum wind speed, and water vapor pressure. The starch and soluble sugar contents in the sapwood of G. floribunda were relatively higher before the growing season. The starch content was lowest in the end of March, while the content of soluble sugars was lowest in middle of May. At the end of the growing season, the contents of starch and soluble sugar in G. floribunda peaked in the middle of October and the end of December, respectively.

Size dependent associations between tree diameter growth rates and functional traits in an Asian tropical seasonal rainforest.

Many studies focus on the relationships between plant functional traits and tree growth performances. However, little is known about the ontogenetic shifts of the relationships between functional traits and tree growth. This study examined associations between stem and leaf functional traits and growth rates and their ontogenetic shifts across 20 tropical tree species in a tropical seasonal rainforest in Xishuangbanna, south-west China. For each species, physiological active branches of individual trees belonged to three size classes (i.e. small, diameter at breast height (DBH) 5-10 cm; middle, DBH 10-20 cm; big, DBH >20 cm) were sampled respectively. We measured 18 morphological and structural traits, which characterised plant hydraulic properties or leaf economic spectrum. Associations between diameter growth rates and functional traits were analysed across three size classes. Our results revealed that diameter growth rates of big-sized trees were mainly related to traits related to plant hydraulic efficiency (i.e. theoretical hydraulic conductivity (Ktheo) and leaf vein density (Dvein)), which suggests that the growth of large trees is limited mainly by their xylem water transport capacity. For middle-sized trees, growth rates were significantly related to traits representing leaf economic spectrum (i.e. specific leaf area (SLA), individual leaf mass (ILM), palisade thickness (PT) and spongy thickness (SP)). Diameter growth rates of small-sized trees were not correlated with hydraulic or leaf economic traits. Thus, the associations between tree growth rates and functional traits are size dependent. Our results suggest ontogenetic shift of functional traits which could potential contribute to different growth response to climate change.

Differential determinants of growth rates in subtropical evergreen and deciduous juvenile trees: carbon gain, hydraulics and nutrient-use efficiencies.

Growth rate varies across plant species and represents an important ecological strategy for competition, resource-use and fitness. However, empirical studies often show a low predictability of functional traits to tree growth. We measured stem diameter and height growth rates (DGRs and HGRs) of 96 juvenile trees (2-5 m tall) of eight evergreen and eight deciduous broadleaf tree species over three consecutive years in a subtropical forest in south-western China. We examined the relationships between tree growth rates and 20 leaf/stem traits that are associated with carbon gain, stem hydraulics and nutrient-use efficiency, as well as the difference between evergreen and deciduous trees. We found that cross-species variations of stem DGR/HGR can be predicted by leaf photosynthetic capacity, leaf mass per area, xylem-theoretical-specific hydraulic conductivity, wood density (WD) and photosynthetic-nutrient-use efficiencies. Higher leaf carbon assimilation and lower leaf/stem constructing costs facilitate deciduous species to be more resource acquisitive and consequently faster growth within a relatively shorter growing season, whereas evergreen species exhibit more conservative strategies and thus slower growth. Furthermore, stem growth rates of evergreen species showed were more dependence on leaf carbon gains, whereas stem hydraulic efficiency was more important for deciduous tree growth. Our results suggest that physiological traits (photosynthesis, hydraulics and nutrient-use efficiency) can predict tree diameter and height growth of subtropical tree species. The differential resource acquisition and use strategies and their associations with tree growth between evergreen and deciduous trees provide insights into explaining the coexistence of evergreen and deciduous tree species in subtropical forests.

Difference between emergent aquatic and terrestrial monocotyledonous herbs in relation to the coordination of leaf stomata with vein traits.

Emergent aquatic plants mostly occur in shallow waters and root in bottom substrates, but their leaves emerge from the water surface and are thus exposed to air, similar to the leaves of terrestrial plants. Previous studies have found coordination between leaf water supply and demand in terrestrial plants; however, whether such a coordination exists in emergent aquatic plants remains unknown. In this study, we analysed leaf veins and stomatal characteristics of 14 emergent aquatic and 13 terrestrial monocotyledonous herb species (EMH and TMH), with 5 EMH and 8 TMH belonging to Poaceae. We found that EMH had significantly higher mean leaf area, leaf thickness, stomatal density, stomatal number per vein length and major vein diameter, but lower mean major vein length per area (VLA) and total VLA than TMH. There was no significant difference in stomatal length, minor VLA and minor vein diameter between the two groups. Stomatal density and total VLA were positively correlated among the EMH, TMH, as well as the 8 Poaceae TMH species, but this correlation became non-significant when data from both the groups were pooled. Our results showed that the differences in water supply between emergent aquatic and terrestrial plants modify the coordination of their leaf veins and stomatal traits.

The contrasting leaf functional traits between a karst forest and a nearby non-karst forest in south-west China.

Karst and non-karst forests occur in the same region in south-west China, but the soil water and mineral nutrients availability are different between the forests. Our hypothesis was that the leaves of karst trees would be better adapted to dry, nutrient-poor conditions than those of trees in a nearby non-karst forest. We compared the gas exchange, anatomical characteristics and mineral nutrient concentrations in leaves from 21 tree species in a tropical karst forest and 19 species in a nearby non-karst forest in south-west China. We found that the leaves of karst trees had higher P concentrations, photosynthetic capacity and water use efficiency, and greater adaxial and abaxial epidermis thickness than leaves of non-karst forest trees. Evergreen and deciduous trees differed more significantly in leaf functional traits in the karst forest than in the non-karst forest. The leaf palisade:spongy mesophyll thickness ratio was positively correlated with stomatal conductance and negatively correlated with photosynthetic water use efficiency in the karst forest but not in the non-karst forest. Our findings indicate that karst forest trees are more conservative in water use, whereas soil P deficiency could be a major limiting factor for the growth of non-karst forest trees.

Leaf turgor loss point is correlated with drought tolerance and leaf carbon economics traits.

Leaf turgor loss point (πtlp) indicates the capacity of a plant to maintain cell turgor pressure during dehydration, which has been proven to be strongly predictive of the plant response to drought. In this study, we compiled a data set of πtlp for 1752 woody plant individuals belonging to 389 species from nine major woody biomes in China, along with reduced sample size of hydraulic and leaf carbon economics data. We aimed to investigate the variation of πtlp across biomes varying in water availability. We also tested two hypotheses: (i) πtlp predicts leaf hydraulic safety margins and (ii) it is correlated with leaf carbon economics traits. Our results showed that there was a positive relationship between πtlp and aridity index: biomes from humid regions had less negative values than those from arid regions. This supports the idea that πtlp may reflect drought tolerance at the scale of woody biomes. As expected, πtlp was significantly positively correlated with leaf hydraulic safety margins that varied significantly across biomes, indicating that this trait may be useful in modelling changes of forest components in response to increasing drought. Moreover, πtlp was correlated with a suite of coordinated hydraulic and economics traits; therefore, it can be used to predict the position of a given species along the 'fast-slow' whole-plant economics spectrum. This study expands our understanding of the biological significance of πtlp not only in drought tolerance, but also in the plant economics spectrum.

Tradeoff between Stem Hydraulic Efficiency and Mechanical Strength Affects Leaf-Stem Allometry in 28 Ficus Tree Species.

Leaf-stem allometry is an important spectrum that linked to biomass allocation and life history strategy in plants, although the determinants and evolutionary significance of leaf-stem allometry remain poorly understood. Leaf and stem architectures - including stem area/mass, petiole area/mass, lamina area/mass, leaf number, specific leaf area (LA), and mass-based leafing intensity (LI) - were measured on the current-year branches for 28 Ficus species growing in a common garden in SW China. The leaf anatomical traits, stem wood density (WD), and stem anatomical and mechanical properties of these species were also measured. We analyzed leaf-stem allometric relationships and their associations with stem hydraulic ad mechanical properties using species-level data and phylogenetically independent contrasts. We found isometric relationship between leaf lamina area/mass and stem area/mass, suggesting that the biomass allocation to leaf was independent to stem size. However, allometric relationship between LA/mass and petiole mass was found, indicating large leaves invest a higher fractional of biomass in petiole than small ones. LI, i.e., leaf numbers per unit of stem mass, was negatively related with leaf and stem size. Species with larger terminal branches tend to have larger vessels and theoretical hydraulic conductivity, but lower WD and mechanical strength. The size of leaf lamina, petiole, and stem was correlated positively with stem theoretical hydraulic conductivity, but negatively with stem WD and mechanical strength. Our results suggest that leaf-stem allometry in Ficus species was shaped by the trade-off between stem hydraulic efficiency and mechanical stability, supporting a functional interpretation of the relationship between leaf and stem dimensions.

Different hydraulic traits of woody plants from tropical forests with contrasting soil water availability.

In southwestern China, tropical karst forests (KF) and non-karst rain forests (NKF) have different species composition and forest structure owing to contrasting soil water availability, but with a few species that occur in both forests. Plant hydraulic traits are important for understanding the species' distribution patterns in these two forest types, but related studies are rare. In this study, we investigated hydraulic conductivity, vulnerability to drought-induced cavitation and wood anatomy of 23 abundant and typical woody species from a KF and a neighboring NKF, as well as two Bauhinia liana species common to both forests. We found that the KF species tended to have higher sapwood density, smaller vessel diameter, lower specific hydraulic conductivity (ks) and leaf to sapwood area ratio, and were more resistant to cavitation than NKF species. Across the 23 species distinctly occurring in either KF or NKF, there was a significant tradeoff between hydraulic efficiency and safety, which might be an underlying mechanism for distributions of these species across the two forests. Interestingly, by possessing rather large and long vessels, the two Bauhinia liana species had extremely high ks but were also high resistance to cavitation (escaping hydraulic tradeoff). This might be partially due to their distinctly dimorphic vessels, but contribute to their wide occurrence in both forests.

Quantifying deforestation and forest degradation with thermal response.

Deforestation and forest degradation cause the deterioration of resources and ecosystem services. However, there are still no operational indicators to measure forest status, especially for forest degradation. In the present study, we analysed the thermal response number (TRN, calculated by daily total net radiation divided by daily temperature range) of 163 sites including mature forest, disturbed forest, planted forest, shrubland, grassland, savanna vegetation and cropland. TRN generally increased with latitude, however the regression of TRN against latitude differed among vegetation types. Mature forests are superior as thermal buffers, and had significantly higher TRN than disturbed and planted forests. There was a clear boundary between TRN of forest and non-forest vegetation (i.e. grassland and savanna) with the exception of shrubland, whose TRN overlapped with that of forest vegetation. We propose to use the TRN of local mature forest as the optimal TRN (TRNopt). A forest with lower than 75% of TRNopt was identified as subjected to significant disturbance, and forests with 66% of TRNopt was the threshold for deforestation within the absolute latitude from 30° to 55°. Our results emphasized the irreplaceable thermal buffer capacity of mature forest. TRN can be used for early warning of deforestation and degradation risk. It is therefore a valuable tool in the effort to protect forests and prevent deforestation.

Corrigendum: Earlywood and Latewood Stable Carbon and Oxygen Isotope Variations in Two Pine Species in Southwestern China during the Recent Decades.

[This corrects the article on p. 2050 in vol. 7, PMID: 28119725.].

Stable stomatal number per minor vein length indicates the coordination between leaf water supply and demand in three leguminous species.

The coordination between minor vein density (MVD) and stomatal density (SD) has been found in many plants. However, we still know little about the influence of leaf node on this correlation relationship. Here, we devised the new functional trait 'stomatal number per minor vein length' (SV). By measuring leaflet area (LA), MVD, SD, and SV, we demonstrated the significance of this functional trait in Arachis hypogaea (peanut) grown under different light regimes and in sun leaves of Dalbergia odorifera and Desmodium renifolium. We found that SV did not change significantly with leaflet node or with LA within each light treatment, while shading caused a significant decrease in SV. The positive correlation between SD and MVD was found in peanut under each light regime. Sun leaves of D. odorifera and D. renifolium also had stable SV along the leaflet node, with a positive correlation between MVD and SD. We conclude that under a certain light regime, a stable SV similar to the positive correlation between MVD and SD can also indicate the coordination between leaf water supply and demand. Our findings highlight the significance of SV and provide new insight into the coordination between stomatal number and minor vein length.

Earlywood and Latewood Stable Carbon and Oxygen Isotope Variations in Two Pine Species in Southwestern China during the Recent Decades.

Stable isotopes in wood cellulose of tree rings provide a high-resolution record of environmental conditions, yet intra-annual analysis of carbon and oxygen isotopes and their associations with physiological responses to seasonal environmental changes are still lacking. We analyzed tree-ring stable carbon (δ13C) and oxygen (δ18O) isotope variations in the earlywood (EW) and latewood (LW) of pines from a secondary forest (Pinus kesiya) and from a natural forest (Pinus armandii) in southwestern China. There was no significant difference between δ13CEW and δ13CLW in P. kesiya, while δ13CEW was significantly higher than δ13CLW in P. armandii. For both P. kesiya and P. armandii, δ13CEW was highly correlated with previous year's δ13CLW, indicating a strong carbon carry-over effect for both pines. The intrinsic water use efficiency (iWUE) in the earlywood of P. armandii was slightly higher than that of P. kesiya, and iWUE of both pine species showed an increasing trend, but at a considerably higher rate in P. kesiya. Respective δ13CEW and δ13CLW series were not correlated between the two pine species and could be influenced by local environmental factors. δ13CEW of P. kesiya was positively correlated with July to September monthly mean temperature (MMT), whereas δ13CEW of P. armandii was positively correlated with February to May MMT. Respective δ18OEW and δ18OLW in P. kesiya were positively correlated with those in P. armandii, indicating a strong common climatic forcing in δ18O for both pine species. δ18OEW of both pine species was negatively correlated with May relative humidity and δ18OEW in P. armandii was negatively correlated with May precipitation, whereas δ18OLW in both pine species was negatively correlated with precipitation during autumn months, showing a high potential for climate reconstruction. Our results reveal slightly higher iWUE in natural forest pine species than in secondary forest pine species, and separating earlywood and latewood of for δ18O analyses could provide seasonally distinct climate signals in southwestern China.

Differences in the responses of photosystem I and photosystem II of three tree species Cleistanthus sumatranus, Celtis philippensis and Pistacia weinmannifolia exposed to a prolonged drought in a tropical limestone forest.

Drought stress can induce closure of stomata, thus leading to photoinhibition. The effects of prolonged severe drought under natural growing conditions on photosystem I (PSI), photosystem II (PSII) and cyclic electron flow (CEF) in drought-tolerant tree species are unclear. In spring 2010, southwestern China confronted severe drought that lasted several months. Using three dominant evergreen species, Cleistanthus sumatranus (Miq.) Muell. Arg. (Euphorbiaceae), Celtis philippensis Bl. (Ulmaceae) and Pistacia weinmannifolia J. Poisson ex Franch. (Anacardiaceae) that are native to a tropical limestone forest, we investigated the influence of this stress on PSI and PSII activities as well as light energy distribution in the PSII and P700 redox state. By the end of the drought period, predawn leaf water potential (Ψ(pd)) largely declined in each species, especially in C. sumatranus. Photosystem I activity strongly decreased in the three species, especially in C. sumatranus which showed a decrease of 65%. The maximum quantum yield of PSII after dark adaptation remained stable in P. weinmannifolia and C. philippensis but significantly decreased in C. sumatranus. Light response curves indicated that both linear electron flow and non-photochemical quenching were severely inhibited in C. sumatranus along with disappearance of CEF, resulting in deleterious excess light energy in PSII. We conclude that PSI is more sensitive than PSII to prolonged severe drought in these three drought-tolerant species, and CEF is essential for photoprotection in them.