Carbonyl sulfide (COS) as a tracer for canopy photosynthesis, transpiration and stomatal conductance: potential and limitations

The theoretical basis for the link between the leaf exchange of carbonyl sulfide (COS), carbon dioxide (CO(2)) and water vapour (H(2)O) and the assumptions that need to be made in order to use COS as a tracer for canopy net photosynthesis, transpiration and stomatal conductance, are reviewed. The ratios of COS to CO(2) and H(2)O deposition velocities used to this end are shown to vary with the ratio of the internal to ambient CO(2) and H(2)O mole fractions and the relative limitations by boundary layer, stomatal and internal conductance for COS. It is suggested that these deposition velocity ratios exhibit considerable variability, a finding that challenges current parameterizations, which treat these as vegetation-specific constants. COS is shown to represent a better tracer for CO(2) than H(2)O. Using COS as a tracer for stomatal conductance is hampered by our present poor understanding of the leaf internal conductance to COS. Estimating canopy level CO(2) and H(2)O fluxes requires disentangling leaf COS exchange from other ecosystem sources/sinks of COS. We conclude that future priorities for COS research should be to improve the quantitative understanding of the variability in the ratios of COS to CO(2) and H(2)O deposition velocities and the controlling factors, and to develop operational methods for disentangling ecosystem COS exchange into contributions by leaves and other sources/sinks. To this end, integrated studies, which concurrently quantify the ecosystem-scale CO(2), H(2)O and COS exchange and the corresponding component fluxes, are urgently needed.     

The potential use of papyrus (Cyperus papyrus L.) wetlands as a source of biomass energy for sub‐Saharan Africa

Four of five people in sub‐Saharan Africa rely on the traditional use of solid biomass, mainly fuelwood, for cooking. In some areas, the current rate of fuelwood consumption will exhaust biomass reserves within the next decade or two. A largely unrecognized source of biomass are tropical wetland ecosystems which have been shown to be some of the most productive ecosystems globally, exhibiting rates of net primary productivity comparable with high‐input, intensively managed agricultural systems. Papyrus (Cyperus papyrus L.) is an emergent sedge with C₄ photosynthesis which is native to the wetlands, river valleys and lakes of central, eastern and southern Africa. The mean standing dry matter of culms and umbels measured at a number of locations throughout East Africa is 38.3 ± 21.6 tDM ha^?1, and the aerial net primary productivity ranges between 25.9 and 136.4 tDM ha^?1 yr^?1. Papyrus vegetation can be harvested by hand and stacked on the rhizome mat for partial air‐drying, and it has been demonstrated that an annual harvesting regime has no negative impacts on long‐term productivity. The use of papyrus as a biofuel for cooking and heating depends on converting it to a suitably combustible form, such as compressed or carbonized briquettes with a calorific value approximately one‐third less than wood charcoal. While papyrus has significant potential as a biofuel, we argue that an integrated management and decision‐making framework for the sustainable utilization of papyrus wetlands is required, in which all ecosystem services including the provision of biomass energy need to be assessed. Sustainability of papyrus wetlands requires management which combines the strength of traditional communal governance and modern legislation to promote its utilization. In this way, local communities can benefit from the inherent advantages of tropical wetlands as very productive ecosystems.     

A coupled model of photosynthesis,stomatal conductance and transpiration for a rose leaf (Rosa hybrida L.)

The following three models were combined to predict simultaneously photosynthesis, stomatal conductance, transpiration and leaf temperature of a rose leaf: the biochemical model of photosynthesis of Farquhar, von Caemmerer and Berry (1980, Planta 149: 78-90), the stomatal conductance model of Ball, Woodrow and Berry (In: Biggens J, ed. Progress in photosynthesis research. The Netherlands: Martinus Nijhoff Publishers), and an energy balance model. The photosynthetic parameters: maximum carboxylation rate, potential rate of electron transport and rate of triose phosphate utilization, and their temperature dependence were determined using gas exchange data of fully expanded, young, sunlit leaves. The stomatal conductance model was calibrated independently. Prediction of net photosynthesis by the coupled model agreed well with the validation data, but the model tended to underestimate rates of stomatal conductance and transpiration. The coupled model developed in this study can be used to assist growers making environmental control decisions in glasshouse production.     

整树水力导度协同冠层气孔导度调节森林蒸腾

冠层气孔导度决定森林的蒸腾效率,它对驱动水汽移动的水汽应力的响应受树木水力结构的影响,并随水汽压亏缺上升和水力导度下降而降低,维持水势在最低阈值之上,避免出现水力灾变,调控冠层蒸腾。由于叶形和树冠结构的特点,部分脱耦联反映了湿润地区阔叶林冠层与大气的水汽交换特征,单纯以气孔导度的变化难以完整描述水分通量的调节规律,因而,需要考虑冠层气孔导度与水力导度协同控制冠层蒸腾的潜在机理。通过整合叶片气孔气体交换、树干液流、冠层微气象和其他环境因子的野外观测值,估测不同时间尺度的森林冠层气孔导度与大气的脱耦联系数和变异范围,以基于树干液流的冠层蒸腾,结合叶片/土壤水势梯度计算的水力导度,分析水力导度影响冠层气孔导度响应水汽压亏缺的敏感性,可以揭示和阐明水力导度和冠层气孔导度联合调节森林蒸腾的机理,对准确估测全球变化背景下森林对水资源利用的潜在生态效应有明显的理论意义。     

Zinc fertilization and water stress affects plant water relations, stomatal conductance and osmotic adjustment in chickpea (Cicer arientinum L.)

Chickpea (Cicer arietinum) is an important dryland pulse crop in many parts of the world. Productivity is often limited by periods of water deficit and in a number of regions zinc deficiency occurs, but the interaction between zinc nutrition and water stress has not been studied extensively. This interaction was examined in two glasshouse experiments. Chickpea was grown under deficient (no applied Zn) or adequate (2.5 μg Zn/g soil) levels of zinc in pots for either 53 days (Experiment 1) or 40 days (Experiment 2) before being exposed to a single period of water stress that lasted for 12 days (Experiment 1) or 23 days (Experiment 2). In one experiment four genotypes (Tyson, ICC-4958, T-1587 and NIFA-88) differing in their sensitivity to zinc deficiency were compared during a single drying cycle, and in the second experiment a single cultivar (Tyson) was compared under well-watered and water stress conditions. Water stress was induced by allowing the soil to dry gradually and the responses in shoot biomass, water use, plant water relations and carbon isotope discrimination (Δ, ‰) were measured. Shoot biomass, water use and water use efficiency were reduced by zinc deficiency. Stomatal conductance was lower in zinc-deficient plants as well. Zinc deficiency reduced Δ by about 1‰ and there were significant differences in Δ between genotypes which were independent of the level of zinc nutrition. At an adequate level of zinc there was a significant negative correlation between Δ and shoot biomass and between Δ and water use efficiency among the four chickpea genotypes, but these correlations were not significant under zinc deficiency. Osmotic potential was lower and turgor higher in the leaves of zinc-deficient plants, but the ability to adjust osmotically was reduced by zinc deficiency as stress developed. In conclusion, zinc-deficiency reduced the efficiency with which the water was used for biomass production and compromised the plant’s capacity to respond to water stress by osmotic adjustment.     

A synthesis of past,current and future research for protection and management of papyrus (Cyperus papyrus L.) wetlands in Africa

Papyrus wetlands (dominated by the giant sedge Cyperus papyrus L.) occur throughout eastern, central and southern Africa and are important for biodiversity, for water quality and quantity regulation and for the livelihoods of millions of people. To draw attention to the importance of papyrus wetlands, a special session entitled “The ecology of livelihoods in papyrus wetlands” was organized at the 9th INTECOL Wetlands Conference in Orlando, Florida in June 2012. Papers from the session, combined with additional contributions, were collected in a special issue of Wetlands Ecology and Management. The current paper reviews ecological and hydrological characteristics of papyrus wetlands, summarizes their ecosystem services and sustainable use, provides an overview of papyrus research to date, and looks at policy development for papyrus wetlands. Based on this review, the paper provides a synthesis of research and policy priorities for papyrus wetlands and introduces the contributions in the special issue. Main conclusions are that (1) there is a need for better estimates of the area covered by papyrus wetlands. Limited evidence suggests that the loss of papyrus wetlands is rapid in some areas; (2) there is a need for a better understanding and modelling of the regulating services of papyrus wetlands to support trade-off analysis and improve economic valuation; (3) research on papyrus wetlands should include assessment of all ecosystem services (provisioning, regulating, habitat, cultural) so that trade-offs can be determined as the basis for sustainable management strategies (‘wise use’); (4) more research on the governance, institutional and socio-economic aspects of papyrus wetlands is needed to assist African governments in dealing with the challenges of conserving wetlands in the face of growing food security needs and climate change. The papers in the special issue address a number of these issues.     

Aquaporin regulation in roots controls plant hydraulic conductance,stomatal conductance,and leaf water potential in Pinus radiata under water stress

Stomatal regulation is crucial for forest species performance and survival on drought‐prone sites. We investigated the regulation of root and shoot hydraulics in three Pinus radiata clones exposed to drought stress and its coordination with stomatal conductance (g_s) and leaf water potential (Ψ_leaf). All clones experienced a substantial decrease in root‐specific root hydraulic conductance (K_root‐r) in response to the water stress, but leaf‐specific shoot hydraulic conductance (K_shoot‐l) did not change in any of the clones. The reduction in K_root‐r caused a decrease in leaf‐specific whole‐plant hydraulic conductance (K_plant‐l). Among clones, the larger the decrease in K_plant‐l, the more stomata closed in response to drought. Rewatering resulted in a quick recovery of K_root‐r and g_s. Our results demonstrated that the reduction in K_plant‐l, attributed to a down regulation of aquaporin activity in roots, was linked to the isohydric stomatal behaviour, resulting in a nearly constant Ψ_leaf as water stress started. We concluded that higher K_plant‐l is associated with water stress resistance by sustaining a less negative Ψ_leaf and delaying stomatal closure.     

Plant biomass and nutrient levels of a tropical macrophyte (Cyperus papyrus L.) receiving domestic wastewater

The aboveground biomass and nutrient content ofCyperus papyrus were determined in a small tropical swamp receiving domestic wastewater. It was found that the biomass (4,955 g.m^–2 dry weight) was the highest ever reported for papyrus. The levels of both nitrogen and phosphorus in the plant organs were very high. The nitrogen concentrations of the various plant organs were 4.8% roots, 8.4% rhizomes, 4.5% scales, 4.8% culms, and 6.2% umbels on dry weight basis. As to phosphorus the concentrations were 0.09% roots, 0.11% rhizomes, 0.09% scales, 0.10% culms, and 0.13% umbels. The high biomass and nutrient contents of the plants may have been caused by the high nutrient levels in the surrounding water. Comparison of the nitrogen to phosphorus ratios in the plants to those in the surrounding water showed that the plants stored very high amounts of nitrogen.     

Nutrient release from swamp water,Cyperus papyrus L. organs and swamp sediments

Nitrogen and phosphorus releases from swamp water,Cyperus papyrus plant organs and papyrus swamp sediments were investigated over five weeks. The release of nutrients by dissolution of decomposing organic matter was very important for the swamp water under the prevailing anaerobic conditions. Large amounts of nutrients were found in the incubation water with plant organs and sediments within the first 48 hours. Leaching of the nutrients from the plant organs and sediment caused high initial values. Subsequently decomposition of the organic material caused slight changes in the nutrient levels in the water. Probably denitrification played a role in the nitrogen dynamics, while phosphorus was mineralized.     

Nutritive value of papyrus (Cyperus papyrus,Cyperaceae), a tropical emergent macrophyte

Papyrus (Cyperus papyrus, Cyperaceae) covers large areas in tropical African wetlands. Analysis of its nutritive value has revealed that crude protein is higher in umbels than culms. Ruminai dry matter digestibility of papyrus is, however, higher in culms than umbels. Both the crude protein and ruminai dry matter digestibility decrease with increasing age of the plant. Values for crude protein and ruminai dry matter digestibility are similar to those reported for the range grasses that constitute the greatest percentage of forage in East Africa. In general, papyrus has some grazing potential and could be used as fodder especially in the dry season when other forage is scarce and of low nutritive value.     

Phreatophytes under stress: transpiration and stomatal conductance of saltcedar (Tamarix spp.) in a high-salinity environment

Background and aims

We sought to understand the environmental constraints on an arid-zone riparian phreatophtye, saltcedar (Tamarix ramosissima and related species and hybrids), growing over a brackish aquifer along the Colorado River in the western U.S. Depth to groundwater, meteorological factors, salinity and soil hydraulic properties were compared at stress and non-stressed sites that differed in salinity of the aquifer, soil properties and water use characteristics, to identify the factors depressing water use at the stress site.

Methods

Saltcedar leaf-level transpiration (E_L), LAI, and stomatal conductance (G_S) were measured over a growing season (June–September) with Granier and stem heat balance sensors and were compared to those for saltcedar at the non-stress site determined in a previous study. Transpiration on a ground-area basis (E_G) was calculated as E_L?×?LAI. Environmental factors were regressed against hourly and daily E_L and G_S at each site to determine the main factors controlling water use at each site.

Results

At the stress site, mean E_G over the summer was only 30 % of potential evapotranspiration (ET_o). G_S and E_G peaked between 8 and 9 am then decreased over the daylight hours. Daytime G_S was negatively correlated with vapor pressure deficit (VPD) (P?<?0.05). By contrast, E_G at the non-stress site tracked the daily radiation curve, was positively correlated with VPD and was nearly equal to ET_o on a daily basis. Depth to groundwater increased over the growing season at both sites and resulted in decreasing E_G but could not explain the difference between sites. Both sites had high soil moisture levels throughout the vadose zone with high calculated unsaturated conductivity. However, salinity in the aquifer and vadose zone was three times higher at the stress site than at the non-stress site and could explain differences in plant E_G and G_S.

Conclusions

Salts accumulated in the vadose zone at both sites so usable water was confined to the saturated capillary fringe above the aquifer. Existence of a saline aquifer imposes several types of constraints on phreatophyte E_G, which need to be considered in models of plant water uptake. The heterogeneous nature of saltcedar E_G over river terraces introduces potential errors into estimates of ET by wide-area methods.     

Epidermal conductance, stomatal density and stomatal size among genotypes of Sorghum bicolor (L.) Moench

Abstract. The ability of a plant to survive severe water deficits depends on its ability to restrict water loss through the leaf epidermis after stomata attain minimum aperture. At this stage, the rate of water loss is regulated by the epidermal conductance (g_c). Low g_c would be a useful selection criterion to identify genotypes with enhanced survival capability. Consequently, variation in g_c among Sorghum bicolor (L.) Moench genotypes was evaluated. Since there is little conclusive evidence linking g _c with leaf waxiness, alternative hypotheses relating g _c to stomatal trails were also examined. Epidermal conductance varied from 6.3 to 17.6mmol m⁻² s⁻¹ among sorghum genotypes. It was unrelated to stomatal pore length which varied with genotype and to pore depth which was similar for all genotypes measured. However, g _c, increased with increasing stomatal density. This indicates that stomatal density plays a direct role in water loss even at very low conductances. The association of low stomatal density with low g _c is consistent with the hypothesis that at the smallest stomata aperture, water loss from the epidermis above guard cell teichodes becomes a significant source of leaf water loss. Since low g _c is directly related to crop survival under severe water deficits, it is recommended that genotypes with low g _c. be selected using the selection criterion of stomatal density.     

A coupled model of stomatal conductance, photosynthesis and transpiration

A model that couples stomatal conductance, photosynthesis, leaf energy balance and transport of water through the soil–plant–atmosphere continuum is presented. Stomatal conductance in the model depends on light, temperature and intercellular CO₂ concentration via photosynthesis and on leaf water potential, which in turn is a function of soil water potential, the rate of water flow through the soil and plant, and on xylem hydraulic resistance. Water transport from soil to roots is simulated through solution of Richards’ equation. The model captures the observed hysteresis in diurnal variations in stomatal conductance, assimilation rate and transpiration for plant canopies. Hysteresis arises because atmospheric demand for water from the leaves typically peaks in mid‐afternoon and because of uneven distribution of soil matric potentials with distance from the roots. Potentials at the root surfaces are lower than in the bulk soil, and once soil water supply starts to limit transpiration, root potentials are substantially less negative in the morning than in the afternoon. This leads to higher stomatal conductances, CO₂ assimilation and transpiration in the morning compared to later in the day. Stomatal conductance is sensitive to soil and plant hydraulic properties and to root length density only after approximately 10 d of soil drying, when supply of water by the soil to the roots becomes limiting. High atmospheric demand causes transpiration rates, LE, to decline at a slightly higher soil water content, θ_s, than at low atmospheric demand, but all curves of LE versus θ_s fall on the same line when soil water supply limits transpiration. Stomatal conductance cannot be modelled in isolation, but must be fully coupled with models of photosynthesis/respiration and the transport of water from soil, through roots, stems and leaves to the atmosphere.     

Environmental and stomatal control of transpiration, canopy conductance and decoupling coefficient in young lemon trees under shading net

Sap flow, as a measure of transpiration, was monitored in 2-year-old lemon trees growing in pots. Eight trees were used in the experiment, four of which were placed under a rectangular shading net, while the other four were maintained in the open air. Daily averages of canopy conductance and photosynthesis were not affected by shading; however, the daily transpiration was reduced in shaded plants, which displayed an increase in water use efficiency compared with exposed trees. The decoupling coefficient was higher in the shaded trees, indicating that the transpiration of lemon trees was efficiently controlled by stomata in exposed plants, while the transpiration rate was mainly influenced by radiation in the plants growing under the net. This influence was more pronounced in the afternoon, when the whole tree transpiration was largely dominated by equilibrium transpiration in the plants under netting, and the relationship between transpiration and radiation showed a steeper slope in shaded trees. The reduction in transpiration and the maintenance of photosynthesis in shaded plants with respect to exposed trees indicated that screen structures in semi-arid and arid environments could be considered as an intermediate solution for reducing plant water stress and increasing water use efficiency.     

Diurnal variations in leaf water potential and stomatal conductance of pigeon pea (Cajanus cajan (L.) Millsp.) cultivars as affected by irrigation levels

Two cultivars of pigeon pea (Cajanus cajan (L.)Millsp.) UPAS-120 and Parbhat were grown in the field under two irrigation treatments: no irrigation and irrigation when cumulative pan evaporation was equal to 50 per cent depletion of available water content in one metre root zone depth. Diurnal changes in leaf water potential and stomatal conductance were recorded on two daysi.e. October 1, 1979 and October 28, 1979 which corresponded to reproductive growth stage of the crop. Plant water potential decreased rapidly during the day up to about 15.00 and increased during evening hours. Higher leaf water potential was recorded in irrigated treatment on both dates. Adaxial and abaxial stomata differed in their response to water stress. Adaxial stomatal conductance started declining during early morning hours, however, abaxial conductance firstly increased up to 09.00 then decreased and increased again in the afternoon except in irrigated crop of cv. UPAS-120 on 28th October, where conductance never increased after 09.00. The reduced rate of stomatal conductance during day hours may be identified as one of the characteristics responsible for drought tolerance in pigeon pea.     

Effects of stand age and tree species on canopy transpiration and average stomatal conductance of boreal forests

We quantified the effect of stand age and tree species composition on canopy transpiration (E_C) by analysing transpiration per unit leaf area (E_L) and canopy stomatal conductance (G_S) for boreal trees comprising a five stand wildfire chronosequence. A total of 196 sap flux sensors were used on 90 trees consisting of Betula papyrifera Marsh (paper birch; present in the youngest stand), Populus tremuloides Michx (quaking aspen), Pinus banksiana Lamb. (jack pine), and Picea mariana (Mill.) (black spruce). While fine roots were positively correlated with stand E_C; leaf area index, basal area, and sapwood area were not. Stands less than 70 years old were dominated by Populus tremuloides and Pinus banksiana and stands greater than 70 years old were composed almost entirely of Picea mariana. As Populus tremuloides and Pinus banksiana increased in size and age, they displayed an increasing sapwood to leaf area ratio (A_S : A_L), a constant minimum leaf water potential (Ψ_L), and a constant proportionality between G_S at low vapour pressure deficit (Dj G_Sref) and the sensitivity of G_S to D (–δ). In contrast, A_S : A_L, minimum Ψ_L, and the proportionally between –δ and G_Sref decreased with height and age in Picea mariana. A G_S model that included the effects of D, A_S : A_L, tree height, and for Picea mariana an increasing soil to leaf water potential gradient with stand age, was able to capture the effects of contrasting hydraulic properties of Picea mariana, Populus tremuloides and Pinus banksiana during stand development after wildfire.     

Relationships between root permeability to water,leaf conductance and transpiration rate in sunflower (Helianthus annuus L.) cultivars

Summary The relations between leaf conductance (g_l) transpiration rate and root permeability to water (R_p) of three sunflower (Helianthus annuus L.) cultivars grown in a controlled environment cabinet are described.No differences in transpiration rates were found but it was shown that plants with low values of R_p have active stomatal closure with favourable consequences for water use efficiency under water limiting conditions.R_p was estimated by applying hydrostatic pressure on the root system. Values of R_p per unit root volume ranged from 0.34×10^–5 to 16.75×10^–5 (s MPa^–1). There were significant inter-cultivar differences (P<0.05) in R_p and g_l and an inverse correlation between R_p and the maximum values cf g_l within cultivars.Pressure applied on the root system is proposed as a useful tool for the determination of differences in the root permeability to water amongst sunflower cultivars.     

Primary productivity of papyrus (Cyperus papyrus) in a tropical swamp; Lake Naivasha,Kenya

Papyrus (Cyperus papyrus) standing biomass and the primary productivity of undisturbed and previously harvested areas of papyrus was measured in Lake Naivasha swamp, Kenya. Papyrus culm density in undisturbed swamp was estimated to be 13·1±1·9 culms m⁻² and aerial biomass was 3602 g m⁻². In undisturbed swamp the aerial productivity was 14·1 g m⁻² day⁻¹ while the previously harvested swamp reached a peak of 21·0 g m⁻² after 6 months. The annual aerial production rate of papyrus in Lake Naivasha was estimated to be 5150 g m⁻² year⁻¹. To sustain yields of regularly harvested papyrus swamps, the harvest intervals should exceed 1 year.     

黄土高原油松冠层气孔导度和蒸腾变化特征与模拟

油松是黄土高原重要的造林树种,模拟其冠层气孔导度和蒸腾对区域水量平衡计算和人工林可持续经营具有重要意义。基于2015—2018年TDP(Thermal dissipation probes)方法所测得液流数据,分析了黄土高原地区油松冠层平均气孔导度(g_c)与冠层蒸腾(Tr)的变化特征与影响因素,并采用Penman-Monteith公式和Jarvis型气孔导度模型模拟了其g_c和Tr的变化过程,结果表明：(1)该地区油松g_c和Tr日内变化均呈现单峰型,日均蒸腾耗水量为(1.25±0.57) mm/d,生长季(4—10月)总蒸腾耗水量均值为195.47 mm。(2)g_c的日内变化受太阳辐射(Rad)驱动(偏相关系数为0.65),当Rad高于300 W/m²时,驱动作用减弱;g_c的日内变化受水汽压亏缺(VPD)控制(偏相关系数为-0.41),随VPD的增加而降低;g_c的日际变化受土壤水分限制(偏相关系数为0.46),当根区相对有效含水率(RE...     

Oscillations in stomatal conductance and plant functioning associated with stomatal conductance: Observations and a model

Summary Measurements of transpiration, leaf water content, and flux of water in a cotton plant exhibiting sustained oscillations, in stomatal conductance are presented, and a model of the mechanism causing this behaviour is developed. The dynamic elements, of the model are capacitors—representing the change of water content with water potential in mesophyll, subsidiary and guard cells—interconnected by resistances representing flow paths in the plant. Increase of water potential in guard cells causes an increase in stomatal conductance. Increase of water potential in the subsidiary cells has the opposite effect and provides the positive feed-back which can cause stomatal conductance to oscillate. The oscillations are shown to have many of the characteristics of free-running oscillations in real plants. The behaviour of the model has been examined, using an analogue computer, with constraints and perturbations representing some of those which could be applied to real plants in physiological experiments. Aspects of behaviour which have been simulated are (a) opening and closing of stomata under the influence of changes in illumination, (b) transient responses due to step changes in potential transpiration, root permeability and potential of water surrounding the roots, (c) the influence of these factors on the occurrence and shape of spontaneous oscillations, and (d) modulation of sustained oscillations due to a circadian rhythm in the permeability of roots.