Comparison of leaf gas exchange and stable isotope signature of water-soluble compounds along canopy gradients of co-occurring Douglas-fir and European beech

Combined δ(13) C and δ(18) O analyses of water-soluble leaf and twig phloem material were used to determine intrinsic water-use efficiency (iWUE) and variability of stomatal conductance at different crown positions in adult European beech (Fagus sylvatica) and Douglas-fir (Pseudotsuga menziesii) trees. Simultaneous gas exchange measurements allowed evaluation of the differences in calculating iWUE from leaf or phloem water-soluble compounds, and comparison with a semi-quantitative dual isotope model to infer variability of net photosynthesis (A(n) ) between the investigated crown positions. Estimates of iWUE from δ(13) C of leaf water-soluble organic matter (WSOM) outperformed the estimates from phloem compounds. In the beech crown, δ(13) C of leaf WSOM coincided clearly with gas exchange measurements. The relationship was not as reliable in the Douglas-fir. The differences in δ(18) O between leaf and phloem material were found to correlate with stomatal conductance. The semi-quantitative model approach was applicable for comparisons of daily average A(n) between different crown positions and trees. Intracanopy gradients were more pronounced in the beech than in the Douglas-fir, which reached higher values of iWUE at the respective positions, particularly under dry air conditions.     

Variability of latewood-widths and -stable isotope ratios in a sessile oak tree (Quercus petraea (Matt.) Liebl.)

Axial variation of latewood-widths and -stable isotopes (δ¹³C and δ¹⁸O) of a single sessile oak tree (Quercus petraea (Matt.) Liebl.) was analyzed. The influence of sampling height was found to be weak for both, latewood-widths and -stable isotopes in the latewood. The observed trends with stem-height show clearly that possible effects due to cambial aging are overrated by annual effects. In addition, relations between samples from different stem-heights are strong for identical calendar years while they are poor for identical cambial age. This indicates that samples from both, living trees and historical wood can be taken from any height position of a trunk or beam as long they are free from anatomical abnormities. Simple correlation and Gleichläufigkeit were used for validation.     

Pleistocene paleoenvironmental evolution at continental middle latitude inferred from carbon and oxygen stable isotope analysis of ostracodes from the Guadix-Baza Basin (Granada, SE Spain)

A representative paleoenvironmental reconstruction of continental middle latitude from ca. 2 my to the upper part of Middle Pleistocene (279 ± 77 ky) was obtained from the carbon and oxygen stable isotopes analyzed in ostracode shells (Cyprideis torosa) recovered in the Guadix-Baza Basin (SE Spain), an intramontaneous closed depression filled by alluvial and lacustrine sediments. This study was performed along a 356-m-thick composite section, dated previously by paleomagnetism and the amino acid racemization method. δ¹³C and δ¹⁸O profiles reflected changes in temperature, the evaporation/infill ratio in the water bodies and the amount of rain. δ¹³C is also affected by changes in plant biomass: periods with high δ¹³C and δ¹⁸O values are associated with warm and dry regimes, and with less vegetation, which, in some cases, coincide with the development of displacive gypsum crystals, whereas low δ¹³C and δ¹⁸O values correlate with cold and humid episodes, which cause more vegetation biomass and, therefore, increasing the input of isotopically light carbon. Intermediate δ¹⁸O values are linked to temperate dry or humid episodes when they coincide with high or low δ¹³C values, respectively. 86 paleoclimatic events were distinguished in the Pleistocene record from the δ¹³C and δ¹⁸O profiles. From both the statistical analysis of the geochemical data and the geological observations, four Cold and Humid Long Periods (low δ¹⁸O) and four Warm and Dry Long Periods (high δ¹⁸O) were defined. This differs with respect to the paleoclimatological behavior established for the Northern Hemisphere where during cold periods (glacial), no water was available while permafrost conditions persisted, whereas in warm episodes (interglacial), higher precipitation rates occurred. Good correspondences between the Guadix-Baza Basin paleoclimatic record and a marine oxygen-isotope sequence, two continental cores and other long Mediterranean paleoenvironmental records (pollen sequences from Israel) were found, which suggested that climate changes in the Guadix-Baza Basin were in tune with global climatic changes.     

Ontogeny and leaf gas exchange mediate the carbon isotopic signature of herbaceous plants

Values (Δ(i)) predicted by a simplified photosynthetic discrimination model, based only on diffusion through air followed by carboxylation, are often used to infer ecological conditions from the 13C signature of plant organs (δ13C(p)). Recent studies showed that additional isotope discrimination (d that includes mesophyll conductance, photorespiration and day respiration, and post-carboxylation discrimination) can strongly affect δ13C(p); however, little is known about its variability during plant ontogeny for different species. Effect of ontogeny on leaf gas exchange rates, Δ(i) , observed discrimination (Δ(p)) and d in leaf, phloem and root of seven herbaceous species at three ontogenetic stages were investigated under controlled conditions. Functional group identity and ontogeny significantly affected Δ(i) and Δ(p). However, predicted Δ(i) did not match Δ(p). d, strongly affected by functional group identity and ontogeny, varied by up to 14 ‰. d scaled tightly with stomatal conductance, suggesting complex controls including changes in mesophyll conductance. The magnitude of the changes in δ13C(p) due to ontogeny was similar to that due to environmental factors reported in other studies. d and ontogeny should, therefore, be considered in ecosystem studies, integrated in ecosystem models using δ13C(p) and limit the applicability of δ13C(leaf) as a proxy for water-use efficiency in herbaceous plants.     

Last‐century changes of alpine grassland water‐use efficiency: a reconstruction through carbon isotope analysis of a time‐series of Capra ibex horns

The ecophysiological response of an alpine grassland to recent climate change and increasing atmospheric CO₂ concentration was investigated with a new strategy to go back in time: using a time‐series of Capra ibex horns as archives of the alpine grasslands' carbon isotope discrimination (¹³Δ). From the collection of the Natural History Museum of Bern, horns of 24 males from the population of the Augstmatthorn–Brienzer Rothorn mountains, Switzerland, were sampled covering the period from 1938 to 2006. Samples were taken from the beginning of each year‐ring of the horns, representing the beginning of the horn growth period, the spring. The horns' carbon ¹³C content (Δ¹³C) declined together with that of atmospheric CO₂ over the 69‐year period, but ¹³Δ increased slightly (+0.4‰), though significantly (P<0.05), over the observation period. Estimated intercellular CO₂ concentration increased (+56 μmol mol^?1) less than the atmospheric CO₂ concentration (+81 μmol mol^?1), so that intrinsic water‐use efficiency increased by 17.8% during the 69‐year period. However, the atmospheric evaporative demand at the site increased by approximately 0.1 kPa between 1955 and 2006, thus counteracting the improvement of intrinsic water‐use efficiency. As a result, instantaneous water‐use efficiency did not change. The observed changes in intrinsic water‐use efficiency were in the same range as those of trees (as reported by others), indicating that leaf‐level control of water‐use efficiency of grassland and forests followed the same principles. This is the first reconstruction of the water‐use efficiency response of a natural grassland ecosystem to last century CO₂ and climatic changes. The results indicate that the alpine grassland community has responded to climate change by improving the physiological control of carbon gain to water loss, following the increases in atmospheric CO₂ and evaporative demand. But, effective leaf‐level water‐use efficiency has remained unchanged.     

Leaf characteristics and gas exchange of the mangrove <Emphasis Type="Italic">Laguncularia racemosa</Emphasis> as affected by salinity

Under constant salinity we analysed the leaf characteristics of Laguncularia racemosa (L.) Gaertn. in combination with gas exchange and carbon isotopic composition to estimate leaf water-use efficiency (WUE) and potential nitrogen-use efficiency (NUE). NaCl was not added to the control plants and the others were maintained at salinities of 15 and 30 ‰ (S₀, S₁₅, and S₃₀, respectively). Leaf succulence, sodium (Na), nitrogen (N), and chlorophyll (Chl) contents increased under salinity. Salinity had a negative impact on net photosynthetic rate (P _N) and stomatal conductance (g _s) at high and moderated irradiances. Potential NUE declined significantly (p<0.05) with salinity by 37 and 58 % at S₁₅ and S₃₀, respectively, compared to S₀ plants. Conversely, compared to S₀ plants, P _N/g _s increased under saline conditions by 12 % (S₁₅) and 50 % (S₃₀). Thus, WUE inferred from P _N/g _s was consistent with salinity improved short-term WUE. Long-term leaf WUE was also enhanced by salinity as suggested by significantly increased leaf δ¹³C with salinity. Improved WUE under salinity explains the eco-physiological success of mangrove species under increasing salinity. Conversely, decline in NUE may pose a problem for L. racemosa under hyper-saline environments regardless of N availability.     

Intrinsic water‐use efficiency of temperate seminatural grassland has increased since 1857: an analysis of carbon isotope discrimination of herbage from the Park Grass Experiment

A 150‐year‐long record of intrinsic water‐use efficiency (W_i) was derived from community‐level carbon isotope discrimination (¹³Δ) in the herbage of the unfertilized, unlimed control treatment (plot 3) of the Park Grass Experiment at Rothamsted (England) between 1857 and 2007. ¹³Δ during spring growth (first cut harvested in June) averaged 21.0‰ (±0.5‰ SD) and has not shown a long‐term trend (P=0.5) since 1857. ¹³Δ of summer/autumn growth (second cut harvested between September and November) increased from 21.3‰ to 22.0‰ (P < 0.001) between 1875 and 2007. W_i during spring growth has therefore increased by 33% since the beginning of the experiment, and W_i of summer/autumn growth has increased by 18%. The variation in ¹³Δ was mainly related to weather conditions. Plant available soil water explained 51% and 40% of the variation in spring growth ¹³Δ and summer/autumn growth ¹³Δ, respectively. In the 1857–2007 period yields have not increased, suggesting that community‐level photosynthesis has not increased either. Therefore, the increased W_i probably resulted from a decreased stomatal conductance. Vapour pressure deficit (VPD) during spring growth (March–June) has not changed since 1915, meaning that instantaneous water‐use efficiency (W_t) in spring time has increased and transpiration has probably decreased, provided that leaf temperature followed air temperature. Conversely, VPD in the months between the first and second cut has increased by 0.07 kPa since 1915, offsetting the effect of increased W_i on W_t during summer and early autumn. Our results suggest that vegetation has adjusted physiologically to elevated CO₂ by decreasing stomatal conductance in this nutrient‐limited grassland.     

Development of Water Stress under Increased Atmospheric CO2 Concentration

The increase in water use efficiency (the ratio of photosynthetic to transpiration rates) is likely to be the commonest positive effect of long-term elevation in CO₂ concentration (CE). This may not necessarily lead to decrease in long-term water use owing to increased leaf area. However, some plant species seem to cope better with drought stress under CE, because increased production of photosynthates might enhance osmotic adjustment and decreased stomatal conductance and transpiration rate under CE enable plants to maintain a higher leaf water potential during drought. In addition, at the same stomatal conductance, internal CO₂ concentration might be higher under CE which results in higher photosynthetic rate. Therefore plants under CE of the future atmosphere will probably survive eventual higher drought stress and some species may even be able to extend their biotope into less favourable sites. This revised version was published online in July 2006 with corrections to the Cover Date.     

Species‐specific stomatal response of trees to drought – a link to vegetation dynamics?

Question: Is stomatal regulation specific for climate and tree species, and does it reveal species‐specific responses to drought? Is there a link to vegetation dynamics? Location: Dry inner alpine valley, Switzerland Methods: Stomatal aperture (θ_E) of Pinus sylvestris, Quercus pubescens, Juniperus communis and Picea abies were continuously estimated by the ratio of measured branch sap flow rates to potential transpiration rates (adapted Penman‐Monteith single leaf approach) at 10‐min intervals over four seasons. Results: θ_E proved to be specific for climate and species and revealed distinctly different drought responses: Pinus stomata close disproportionately more than neighbouring species under dry conditions, but has a higher θ_E than the other species when weather was relatively wet and cool. Quercus keeps stomata more open under drought stress but has a lower θ_E under humid conditions. Juniperus was most drought‐tolerant, whereas Picea stomata close almost completely during summer. Conclusions: The distinct microclimatic preferences of the four tree species in terms of θ_E strongly suggest that climate (change) is altering tree physiological performances and thus species‐specific competitiveness. Picea and Pinus currently live at the physiological limit of their ability to withstand increasing temperature and drought intensities at the sites investigated, whereas Quercus and Juniperus perform distinctly better. This corresponds, at least partially, with regional vegetation dynamics: Pinus has strongly declined, whereas Quercus has significantly increased in abundance in the past 30 years. We conclude that θ_E provides an indication of a species' ability to cope with current and predicted climate.     

Detection of breastfeeding and weaning in modern human infants with carbon and nitrogen stable isotope ratios

Carbon ((13)C/(12)C) and nitrogen ((15)N/(14)N) stable isotope ratios were longitudinally measured in fingernail and hair samples from mother-infant pairs where infants were exclusively breastfed (n = 5), breast- and formula-fed (n = 2), or exclusively formula-fed (n = 1) from birth. All exclusively breastfed infants had a dual enrichment in carbon ( approximately 1 per thousand) and nitrogen ( approximately 2-3 per thousand) when compared to maternal values. In contrast, breast- and formula-fed subjects had reduced enrichments compared to exclusively breastfed subjects, and the exclusively formula-fed infant showed no increase in delta(13)C or delta(15)N values. This finding of a carbon trophic level effect in breastfeeding infants suggests that (13)C-enrichments of approximately 1 per thousand in archaeological populations are not necessarily the result of the consumption of C(4)-based weaning foods such as maize or millet. During the weaning process, the delta(13)C results for breastfed infants declined to maternal levels more rapidly than the delta(15)N results. This suggests that delta(13)C values have the potential to track the introduction of solid foods into the diet, whereas delta(15)N values monitor the length of time of breast milk consumption. These findings can be used to refine the isotopic analysis of breastfeeding and weaning patterns in past and modern populations.     

黄龙山林区不同演替时期典型树种光诱导的气孔动力学研究

光诱导的气孔动力学响应快慢是影响植物叶水分利用效率的重要因素,为探索黄土高原不同演替阶段树种水分利用效率差异的生理机制,该研究以黄龙山林区典型树种(演替早期种山杨和白桦、演替后期种辽东栎)的幼龄实生苗为材料,采用盆栽试验,研究了叶片光诱导的气孔导度动力学参数差异及其与气孔特征、叶长期水分利用效率的关系。结果表明：(1)山杨和白桦气孔开放过程中气孔导度(g_s)增加的时间常数(K_i)小于辽东栎,但气孔关闭过程中气孔导度降低的时间常数(K_d)则大于辽东栎,表明山杨和白桦气孔开放更快,而辽东栎的气孔关闭更快。同时,气孔开放过程中山杨和白桦的g_s响应幅度均大于辽东栎,气孔关闭过程中山杨的g_s响应幅度亦大于辽东栎。(2)3种树种中,辽东栎的气孔密度最大,气孔最小,气孔指数最大,辽东栎气孔特征无法解释其慢速的气孔开放过程。 (3)山杨和白桦具有高的光合速率、最大羧化效率和最大电子传递速率,3种树种碳同位素比率(δ¹³C)表征的长期水分利用效率表现为山杨>白桦>辽东栎。研究认为,演替早期种山杨和白桦的高水分利用效率与其快速的气孔开放有关,而演替后期种辽东栎快速的气孔关闭并未增加其水分利用效率,且长期水分利用效率低于山杨和白桦,可能与辽东栎慢速的气孔开放限制了其光合速率有关。     

Diet and dental caries among later stone age inhabitants of the Cape Province, South Africa.

Stable carbon isotope measurements and incidences of dental caries are presented for three groups of prehistoric human skeletons from different regions of the Cape Province, South Africa. The isotopic analyses of bone collagen demonstrate the importance of marine foods in the diet and vary through time, across space, and according to sex. The incidence of dental caries ranges from 0% among heavily marine-dependent individuals from the south-western Cape coast, to 17.7% among skeletons from an archaeological site on the south coast. The extremely high incidence of caries in a hunter-gatherer population may be related to lack of fluoride in the water.