Seasonal Variation of δ^13C of Four Tree Species： A Biological Integrator of Environmental Variables

Foliar δ^13C values, an indicator of long-term intercellular carbon dioxide concentration and, thus, of long-term water use efficiency （WUE） in plants, were measured for Pinus massoniana Lamb., P. elliottii Engelm., Cunninghamia laceolata （Lamb.） Hook., and Schima superba Gardn. et Champ. in a restored forest ecosystem in the Jiazhu River Basin. Seasonal variation and the relationship between the foliar δ^13C values of the four species and environmental factors （monthly total precipitation, monthly average air temperature, relative humidity, atmospheric pressure, and monthly total solar radiation and evaporation） were investigated. The monthly δ^13C values and WUE of the four species increased with increasing precipitation, air temperature, solar radiation, and evaporation, whereas δ^13C values of the four species decreased with increasing relative humidity and atmospheric pressure. Despite significant differences in δ^13C seasonal means for the four species, our results demonstrate a significant convergence in the responses of δ^13C values and WUE to seasonal variations in environmental factors among the species investigated and that the δ^13C signature for each species gives a strong indication of environmental variables.     

Correlations Between Foliar Stable Carbon Isotope Composition and Environ-mental Factors in Desert Plant Reaumuria soongorica (Pall.) Maxim.

Leaves of 407 individuals of Reaumuria soongorica (Pall.) Maxim. collected from the major distribution areas were measured to investigate the distribution characteristics of the stable carbon isotope in this desert plant, as well as correlations between δ13C values and environmental factors. Results showed that δ13C values in R. soongorica ranged from-22.77‰. to-29.85‰. and that the mean δ13C value (-26.52‰)was higher than a previously reported δ13C value for a different desert ecosystem. This indicates that R.soongorica belongs to the C3 photosynthetic pathway and has higher water use efficiency than other species. The correlations between δ13C values and environmental factors demonstrated that the foliar δ13C values in R. soongorica increased significantly with decreasing mean annual precipitation and mean relative humidity, and decreased with decreasing duration of sunshine and evaporation. The spatial distribution trend of δ13C values in R. soongorica was not obvious and there was no significant correlation between the δ13C values and mean annual temperature. We conclude that different distribution trends in δ13C values for R. soongorica were likely caused by stomatal limitation rather than by nutrient-related changes in photosynthetic efficiency and that precipitation played an important role in the wide distribution range of R.soongorica. This pattern of δ13C values for R. soongorica reinforced that it is a super-xerophil in terms of its adaptive strategies to a desert environment.     

Correlations Between Foliar Stable Carbon Isotope Composition and Environmental Factors in Desert Plant Reaumuria soongorica （Pall.） Maxim.

Leaves of 407 individuals of Reaumuria soongorica （Pall.） Maxim. collected from the major distribution areas were measured to investigate the distribution characteristics of the stable carbon isotope in this desert plant, as well as correlations between δ^13C values and environmental factors. Results showed that δ^13C values in R. soongorica ranged from -22.77‰ to -29.85‰ and that the mean δ^13C value （-26.52‰） was higher than a previously reported δ^13C value for a different desert ecosystem. This indicates that R. soongorica belongs to the C3 photosynthetic pathway and has higher water use efficiency than other species. The correlations between δ^13C values and environmental factors demonstrated that the foliar δ^13C values in R. soongorica increased significantly with decreasing mean annual precipitation and mean relative humidity, and decreased with decreasing duration of sunshine and evaporation. The spatial distribution trend of δ^13C values in R. soongorica was not obvious and there was no significant correlation between the δ^13C values and mean annual temperature. We conclude that different distribution trends in δ^13C values for R. soongorica were likely caused by stomatal limitation rather than by nutrient-related changes in photosynthetic efficiency and that precipitation played an important role in the wide distribution range of R. soongorica. This pattern of δ^13C values for R. soongorica reinforced that it is a super-xerophil in terms of its adaptive strategies to a desert environment.     

Stipa tenacissima Does not Affect the Foliar δ13C and δ15N of Introduced Shrub Seedlings in a Mediterranean Semi-arid Steppe

Recent studies have shown that the tussock grass Stipa tenacissima L. facilitates the establishment of late-successional shrubs, in what constitutes the first documented case of facilitation of woody plants by grasses. With the aim of increasing our knowledge of this interaction, in the present study we investigated the effects of S. tenacissima on the foliar δ13C, δ15N, nitrogen concentration, and carbon ： nitrogen ratio of introduced seedlings of Pistacia lentiscus L., Quercus coccifera L., and Medicago arborea L. in a semi-arid Mediterranean steppe. Six months after planting, the values of δ13C ranged between -26.9‰ and -29.6‰, whereas those of δ15N ranged between -1.9‰ and 2.7‰. The foliar C ： N ratio ranged between 10.7 and 53.5, and the nitrogen concentration ranged between 1.0% and 4.4%. We found no significant effect of the microsite provided by S. tenacissima on these variables in any of the species evaluated. The values of δ13C were negatively correlated with predawn water potentials in M. arborea and were positively correlated with relative growth rate in Q. coccifera. The values of δ15N were positively correlated with the biomass allocation to roots in the latter species. The present results suggest that the modification of environmental conditions in the are surrounding S. tenacissima was not strong enough to modify the foliar isotopic and nitrogen concentration of shrubs during the early stages after planting.     

Water sources of dominant species in three alpine ecosystems on the Tibetan Plateau, China

Plant water sources were estimated by two or three compartment linear mixing models using hydrogen and oxygen isotope （δD and δ^18O） values of different components such as plant xylem water, precipitation and river water as well as soil water on the Tibetan Plateau in the summer of 2005. Four dominant species （Quercus aquifolioides, Pinus tabulaeformis, Salix rehderiana and Nitraria tangutorum） in three typical ecosystems （forest, shrub and desert） were investigated in this study. Stable isotope ratios of the summer precipitations and the soil water presented variations in spatial and temporal scales. δ^18O values of N. tangutorum xylem water were constant in the whole growth season and very similar to those of deep soil water. Water sources for all of the plants came from both precipitations and soil water. Plants switched rapidly among different water sources when environmental water conditions changed. Rainwater had different contributions to the plants, which was influenced by amounts of precipitation. The percentage of plant xylem water derived from rainwater rose with an increase in precipitation. Water sources for broad-leaved and coniferous species were different although they grew in the same environmental conditions. For example, the broad-leaved species Q. aquifolioides used mainly the water from deep soil, while 92.5% of xylem water of the coniferous species P. tabulaeformis was derived from rainwater during the growth season. The study will be helpful for us to fully understand responses of species on the Tibetan Plateau to changes in precipitation patterns, and to assess accurately changes of vegetation distribution in the future.     

The changes in water-use efficiency and stoma density of Leymus chinensis along Northeast China Transect

Leymus chinensis is a keystone species in the temperate zone grassland of China. Along the NECT (Northeast China Transect) in 2001, water-use efficiency of L. chinensis was analyzed with δ¹³C, and changes in the stoma density of its leaves were observed and computed under a microscope. Results showed that the ecological plasticity modulation of the stoma density of L. chinensis and its water-use efficiency were two important mechanisms for its broad ecological adaptability. From east to west along the NECT, the δ¹³C of the species varied from -27.49‰ to -23.57‰, consisting with the reduction of annual precipitation, soil water and annual average temperature, but increased with the increase of the elevation of sampling sites. The stoma density increased from 96.9/mm² to 169.5/mm², indicating that the water-use efficiency for the species was improved along the gradient. The linear coefficient between the two parameters was significant (R² = 0.7338). The results of a stepwise regression analysis showed that the soil water was the first marked factor for determining the stoma density, and the next was the annual precipitation, which suggested that the water factors were the primary ecological factors influencing the stoma density of L. chinensis. The findings in this study showed that the responses of the stoma density and the water-use efficiency for L. chinensis to environmental changes were very complicated. They may be the outcome operated synthetically by all environmental factors in the long-term adaptation to different ecological environments, including human activity, for L. chinensis.     

The changes in water-use efficiency and stoma density of Leymus chinensis along Northeast China Transect

Leymus chinensis is a keystone species in the temperate zone grassland of China. Along the NECT (Northeast China Transect) in 2001, water-use efficiency of L. chinensis was analyzed with δ¹³C, and changes in the stoma density of its leaves were observed and computed under a microscope. Results showed that the ecological plasticity modulation of the stoma density of L. chinensis and its water-use efficiency were two important mechanisms for its broad ecological adaptability. From east to west along the NECT, the δ¹³C of the species varied from -27.49‰ to -23.57‰, consisting with the reduction of annual precipitation, soil water and annual average temperature, but increased with the increase of the elevation of sampling sites. The stoma density increased from 96.9/mm² to 169.5/mm², indicating that the water-use efficiency for the species was improved along the gradient. The linear coefficient between the two parameters was significant (R² = 0.7338). The results of a stepwise regression analysis showed that the soil water was the first marked factor for determining the stoma density, and the next was the annual precipitation, which suggested that the water factors were the primary ecological factors influencing the stoma density of L. chinensis. The findings in this study showed that the responses of the stoma density and the water-use efficiency for L. chinensis to environmental changes were very complicated. They may be the outcome operated synthetically by all environmental factors in the long-term adaptation to different ecological environments, including human activity, for L. chinensis.     

鼎湖山马尾松径向生长动态与气候因子的关系

Dendroclimatology method was used to study the relationship between ring index of Pinus massoniana and three climate factors in Dinghushan,South China.The ring indices were negatively correlated with the average tem-perature in June,August and September,and positively correlated with the average temperature of March,whereas no statistically significant correlation was found with monthly precipitation,showing that high summer temperature could limit the radial growth of the species,Meanwhile,tree ring indices showed a strong positive correlation with the relative humidity of April June,August,September,October and November,and also with the average humidity of the whole year,indicating that atmospheric moisture could not fully satisfy the growth of Pinus massoniana despite of abundant rainfall,The possible impact of climate change on the growth of Pinus massoniana was discussed accordingly.     

Seasonal variation and correlation with environmental factors of photosynthesis and water use efficiency of Juglans regia and Ziziphus jujuba

Both the photosynthetic light curves and CO2 curves of Juglans regia L. and Ziziphus jujuba Mill. var. spinosa in three seasons were measured using a LI-6400 portable photosynthesis system. The maximal net photosynthetic rate （Amax）, apparent quantum efficiency（φ）, maximal carboxylation rate （Vcmax） and water use efficiency （WUE） of the two species were calculated based on the curves. The results showed that Amax of J. regia reached its maximum at the late-season, while the highest values of Amax of Z. jujuba occurred at the mid-season. The Amax of J. regia was more affected by relative humidity （RH） of the atmosphere, while that of Z. jujuba was more affected by the air temperature. Light saturation point （LSP） and Light compensation point （LCP） of J. regia had a higher correlation with RH of the atmosphere, those of Z. jujuba, however, had a higher correlation with air temperature. Vcmax of both J. regia and Z. jujuba had negative correlation with RH of the atmosphere. WUE of J. regia would decrease with the rise of the air temperature while that of Z. jujuba increased. Thus it could be seen that RH, temperature and soil moisture had main effect on photosynthesis and WUE of J. regia and Z. jujuba. Incorporating data on the physiological differences among tree species into forest carbon models will greatly improve our ability to predict alterations to the forest carbon budgets under various environmental scenarios such as global climate change, or with differing species composition.     

Patterns of δ^13C and δ^15N in wolverine Gulo gulo tissues from the Brooks Range,Alaska

Knowledge of carnivore diets is essential to understand how carnivore populations respond demographically to variations in prey abundance. Analysis of stable isotopes is a useful complement to traditional methods of analyzing carnivore diets. We used data on δ^13C and δ^15N in wolverine tissues to investigate patterns of seasonal and annual diet variation in a wolverine Gulo gulo population in the western Brooks Range, Alaska, USA. The stable isotope ratios in wolverine tissues generally reflected that of terrestrial carnivores, corroborating previous diet studies on wolverines. We also found variation in δ^13C and δ^15N both between muscle samples collected over several years and between tissues with different assimilation rates, even after correcting for isotopic fractionation. This suggests both annual and seasonal diet variation. Our results indicate that data on δ^13C and δ^15N holds promise for qualitative assessments of wolverine diet changes over time. Such temporal variation may be important indicators of ecological responses to environmental perturbations, and we suggest that more refined studies of stable isotopes may be an important tool when studying temporal change in diets of wolverines and similar carnivores [ Current Zoology 55 （3）： 188- 192, 2009].     

Effect of Seasonality on Distribution of Macroalgae in a Stream System （Xin＇an Spring）in Shanxi Province, North China

A survey of the seasonal distribution of macroalgae in a stream system in Shanxi Province, north China, was undertaken from July 2004 to April 2005. The relative abundance and percentage cover of macroalgae, and several environmental factors were monitored along a 20-m stretch at each of four sites, at intervals of approximately three months （one sampling per season）. Several stream conditions were relatively constant over the sampling period （pH, maximum width and maximum depth）, whereas others exhibited a distinct seasonal pattern （water temperature and specific conductance）, and some fluctuated with no discernable seasonal pattern （current velocity and dissolved oxygen）. Forty-two species of macroalgae were found, with a predominance of Chlorophyta （26 species, 61.9%）. Rhodophyts and Charophyta represented the smallest proportion （1 species each, 2.38%）. Six macroalgae species were the most widespread, occurring in all four sampling sites. Twelve species were found at only one site each. In terms of seasonality, eight species occurred throughout the year, whereas 16 species were found in only one season each. The macroalgal community at Xin＇an Spring was species rich relative to other streams. Species richness per sampling site was negatively correlated with pH. Principal component analysis revealed that no single variable had much influence on the macroalgal seasonal dynamics. We calculated Sorensen similarity indices to compare our study with other continent-wide surveys of stream macroalgae, but the similarity indices were all very low. This study also shows that macroalgae in different locations have significant reproductive isolation.     

Humidity and seasonality drives body size patterns in males of the bush cricket Isophya rizeensis Sevgili, 2003 （Orthoptera： Tettigoniidae： Phaneropterinae）

Two primary patterns of body size variation have been recorded in ectotherms in relation to latitudinal/altitudinal shifts. In some, body size increases with increasing latitude/altitude whereas, in others, body size decreases with increasing latitude/altitude. This clinal variation is generally assumed to be caused by local adaptation to environmental conditions however the selective variable（s） （temperature, humidity, diet quality, etc.） is still heavily debated. Here we investigate geographic variation in body size of dark and pale color morphs of males of the bush-cricket lsophya rizeensis collected from 15 locations along an elevation gradient ranging from 350 to 2 500 m. Using an information theoretical approach we evaluate the relative support of four different hypotheses （the temperature size rule, the moisture gradient hypothesis, the seasonal constraint hypothesis, and the primary productivity hypothesis） explaining body size variation along the altitudinal gradient. Body size variation in pale color morphs showed a curvilinear relationship with altitude while dark color morphs showed no variation in body size. Body size variation in pale color morphs was highly correlated with precipitation and temperature seasonality values thus giving strong support for the moisture gradient and seasonal constraint hypothesis. Our results reinforce the importance of gradients in humidity and seasonality over temperature in the creation of altitudinal body size clines and the role of selection for resistance to stress factors in the establishment of these clines. Whether a body size cline is observed or not might also depend on the phenotypic properties of the individuals, like coloration.     

Estimation of evaporation rate from soil surface using stable isotopic composition of throughfall and stream water in a tropical seasonal rain forest of Xishuangbanna, Southwest China

Xishuangbanna is located at the northern edge of the distribution of tropical forests in Southeast Asia, and it has a very high frequency of radiation fog, especially during the dry season (November-April). In this study, rainwater, throughfall, intercepted fog water (fog drip) by forest canopy, and stream water were collected in January 2002 and December 2003 for stable isotopic analysis at a tropical seasonal rain forest site in Xishuangbanna, Southwest China. The objective of the study is to estimate the evaporation rate from the forest floor. The stable hydrogen (δD) and oxygen isotope composition (δ¹⁸O) of rainwater, throughfall, fog drip, and stream water was determined using an isotope ratio mass spectrometer. Stream water during the non-storm runoff period was considered to reflect the effect of evaporation from the forest floor. The evaporation rates from the forest floor were estimated using isotope composition values in stream water and the total throughfall was estimated using the Rayleigh distillation equation under equilibrium conditions. The results indicated that the annual weighted means of δD and δ¹⁸O in fog drip were consistently more enriched than those of rainwater and stream water, and the fog drip was thought to contain water that had evaporated and recycled. The weighted means of δD and δ¹⁸O in stream water during the non-storm runoff period were 5.69% and 0.39% more enriched than those of the total throughfall (rain throughfall fog drip) in 2002, while in 2003 these were 5.05% and 0.28%, respectively. Evapotranspiration in the humid year 2002 and the dry year 2003, computed from the water balance, was 1186 mm and 987 mm, respectively, which was quite low when compared with the values reported in some humid tropical forests. Consequently, the evaporation rate from the forest floor was 5.1% of the evapotranspiration in 2002, and 3.1% in 2003. The lower evaporation rate was thought to be mainly a result of the influence of the high frequency of heavy radiation fog on the rain forest during the night in the dry season (November-April).     

Estimation of evaporation rate from soil surface using stable isotopic composition of throughfall and stream water in a tropical seasonal rain forest of Xishuangbanna, Southwest China

Xishuangbanna is located at the northern edge of the distribution of tropical forests in Southeast Asia, and it has a very high frequency of radiation fog, especially during the dry season (November-April). In this study, rainwater, throughfall, intercepted fog water (fog drip) by forest canopy, and stream water were collected in January 2002 and December 2003 for stable isotopic analysis at a tropical seasonal rain forest site in Xishuangbanna, Southwest China. The objective of the study is to estimate the evaporation rate from the forest floor. The stable hydrogen (δD) and oxygen isotope composition (δ¹⁸O) of rainwater, throughfall, fog drip, and stream water was determined using an isotope ratio mass spectrometer. Stream water during the non-storm runoff period was considered to reflect the effect of evaporation from the forest floor. The evaporation rates from the forest floor were estimated using isotope composition values in stream water and the total throughfall was estimated using the Rayleigh distillation equation under equilibrium conditions. The results indicated that the annual weighted means of δD and δ¹⁸O in fog drip were consistently more enriched than those of rainwater and stream water, and the fog drip was thought to contain water that had evaporated and recycled. The weighted means of δD and δ¹⁸O in stream water during the non-storm runoff period were 5.69% and 0.39% more enriched than those of the total throughfall (rain throughfall + fog drip) in 2002, while in 2003 these were 5.05% and 0.28%, respectively. Evapotranspiration in the humid year 2002 and the dry year 2003, computed from the water balance, was 1186 mm and 987 mm, respectively, which was quite low when compared with the values reported in some humid tropical forests. Consequently, the evaporation rate from the forest floor was 5.1% of the evapotranspiration in 2002, and 3.1% in 2003. The lower evaporation rate was thought to be mainly a result of the influence of the high frequency of heavy radiation fog on the rain forest during the night in the dry season (November-April).     

Aggregation and mating success of Capnodis tenebrionis （Coleoptera： Buprestidae）

An understanding of the relative importance of extrinsic and intrinsic factors in determining the potential distribution and mating success of individuals is critical for the successful monitoring and management of pest species. Using a combination of field observations and a caged field experiment, we explored the roles of environmental and individual variation on the formation of mating aggregations and mating success in the buprestid beetle Capnodis tenebrionis （Linnaeus, 1767）, a pest species of stone fruit trees. Our field observations revealed that the formation of aggregations is influenced by a range of environmental factors including temperature, photoperiod, and population density. However, aggregations were not at random and were more likely to occur on the section of the plant with highest incidence of solar radiation and thus higher temperatures. Data from our experiment with caged beetles in the field further indicate that the reproductive behavior of this species varies with temperature. The probability of a successful mating occurring was also positively related to both male and female size. Females of C. tenebrionis mate several times over a 4-h period, but generally not with the same male. Information obtained from these studies is useful to define the most appropriate time for pest control, especially adopting strategies that interfere with reproduction.     

Drought-resistant physiological characteristics of four shrub species in arid valley of Minjiang River, China

The formation of 161 km-long arid valley (170 km²) of Mingjiang River is attributed to the effect of foehn. The arid valley presents harsh conditions for the growth of vegetation, which include higher annual evaporation and transpiration rate (1400–2000 mm), lower annual precipitation rate (400–700 mm), steep slope, and infertile soils. Although the vegetation is dominated by xerophilous bushes and grasses, it plays a significant role in soil and water conservation in watershed and provides important references in the arid valley as it is very difficult to grow trees. Drought-resistant physiological characteristics of four native species, i.e., Sophora davidiana, Bauhinia faberi var. microphylla, Convolvulus tragacanthoides, and Artemisia gmelinii, have been measured. It was found that the content of chlorophyll, free proline and soluble sugars showed an increase, whereas evaporation ratio, WSD, RWC and cytolemma osmosis showed a decrease under drought stress. Integrated assessment of the four species indicated that the integrated drought resistance index was the highest in July (0.507), followed by September, August and June, whereas it was the lowest in May (0.442). The result provided a basis for vegetation restoration, especially it improved survival of planted species in May and June when plant species exhibit the weakest drought resistance.     

Carbon Isotope Discrimination is not Correlated with Transpiration Efficiency in Three Cool-Season Grain Legumes （Pulses）

The carbon isotope discrimination （δ^13C） of leaves has been shown to be correlated with the transpiration efficiency of leaves in a wide range of species. This has led to δ^13C being used in breeding programs to select for improved transpiration efficiency. The correlation between δ^13C and transpiration efficiency was determined under well-watered conditions during the vegetative phase in six genotypes of lentil （Lens culinaris Medikus）, six genotypes of chickpea （Cicerarietinum L.） and 10 cultivars of narrow-leafed lupin （Lupinus angustifolius L.）. Biomass （dry matter） accumulation and water use （transpiration） varied among the genotypes in all three species and transpiration efficiency was 40% to 75% higher in the most efficient compared with the least efficient genotypes. However, δ^13C and transpiration efficiency were not significantly correlated in any of the species. This suggests that the δ^13C technique cannot be used in selection for transpiration efficiency in the three grain legumes （pulses） studied.     

Drought-resistant physiological characteristics of four shrub species in arid valley of Minjiang River, China

The formation of 161 km-long arid valley (170 km²) of Mingjiang River is attributed to the effect of foehn. The arid valley presents harsh conditions for the growth of vegetation, which include higher annual evaporation and transpiration rate (1400–2000 mm), lower annual precipitation rate (400–700 mm), steep slope, and infertile soils. Although the vegetation is dominated by xerophilous bushes and grasses, it plays a significant role in soil and water conservation in watershed and provides important references in the arid valley as it is very difficult to grow trees. Drought-resistant physiological characteristics of four native species, i.e., Sophora davidiana, Bauhinia faberi var. microphylla, Convolvulus tragacanthoides, and Artemisia gmelinii, have been measured. It was found that the content of chlorophyll, free proline and soluble sugars showed an increase, whereas evaporation ratio, WSD, RWC and cytolemma osmosis showed a decrease under drought stress. Integrated assessment of the four species indicated that the integrated drought resistance index was the highest in July (0.507), followed by September, August and June, whereas it was the lowest in May (0.442). The result provided a basis for vegetation restoration, especially it improved survival of planted species in May and June when plant species exhibit the weakest drought resistance.