Effects of N fertilization on forest trees and ecosystems: water use, WUE, growth

During my three academic years, I focused on the effects of N fertilization on growth and function of plants and forest stands. The study had the dual objective of estimating the effects of atmospheric N deposition and evaluating the potential management value of N fertilization itself. In particular, the analysis took into account the changes induced in water use and intrinsic transpiration efficiency (ITE), an aspect often overlooked in world literature but of great importance especially in Mediterranean environment, where the positive effects of N fertilization may be denied by the parallel increased transpiration and exacerbated water stress.

Stretching water - Queensland’s water use efficiency cotton and grains adoption program

The Cotton and Grain Adoption Program of the Queensland Rural Water Use Efficiency Initiative is targeting five major irrigation regions in the state with the objective to develop better irrigation water use efficiency (WUE) through the adoption of best management practices in irrigation. The major beneficiaries of the program will be industries, irrigators and local communities. The benefits will flow via two avenues: increased production and profit resulting from improved WUE and improved environmental health as a consequence of greatly reduced runoff of irrigation tailwater into rivers and streams. This in turn will reduce the risk of nutrient and pesticide contamination of waterways. As a side effect, the work is likely to contribute to an improved public image of the cotton and grain industries. In each of the five regions, WUE officers have established grower groups to assist in providing local input into the specific objectives of extension and demonstration activities. The groups also assist in developing growers' perceptions of ownership of the work. Activities are based around four on-farm demonstration sites in each region where irrigation management techniques and hardware are showcased. A key theme of the program is monitoring water use. This is applied both to on-farm storage and distribution as well as to application methods and in-field management. This paper describes the project, its activities and successes.

Effect of irrigation amounts applied with subsurface drip irrigation on corn evapotranspiration, yield, water use efficiency, and dry matter production in a semiarid climate

Quantifying the local crop response to irrigation is important for establishing adequate irrigation management strategies. This study evaluated the effect of irrigation applied with subsurface drip irrigation on field corn (Zea mays L.) evapotranspiration (ETc), yield, water use efficiencies (WUE = yield/ETc, and IWUE = yield/irrigation), and dry matter production in the semiarid climate of west central Nebraska. Eight treatments were imposed with irrigation amounts ranging from 53 to 356 mm in 2005 and from 22 to 226 mm in 2006. A soil water balance approach (based on FAO-56) was used to estimate daily soil water and ETc. Treatments resulted in seasonal ETc of 580-663 mm and 466-656 mm in 2005 and 2006, respectively. Yields among treatments differed by as much as 22% in 2005 and 52% in 2006. In both seasons, irrigation significantly affected yields, which increased with irrigation up to a point where irrigation became excessive. Distinct relationships were obtained each season. Yields increased linearly with seasonal ETc (R 2 = 0.89) and ETc/ETp (R 2 = 0.87) (ETp = ETc with no water stress). The yield response factor (ky), which indicates the relative reduction in yield to relative reduction in ETc, averaged 1.58 over the two seasons. WUE increased non-linearly with seasonal ETc and with yield. WUE was more sensitive to irrigation during the drier 2006 season, compared with 2005. Both seasons, IWUE decreased sharply with irrigation. Irrigation significantly affected dry matter production and partitioning into the different plant components (grain, cob, and stover). On average, the grain accounted for the majority of the above-ground plant dry mass (≈59%), followed by the stover (≈33%) and the cob (≈8%). The dry mass of the plant and that of each plant component tended to increase with seasonal ETc. The good relationships obtained in the study between crop performance indicators and seasonal ETc demonstrate that accurate estimates of ETc on a daily and seasonal basis can be valuable for making tactical in-season irrigation management decisions and for strategic irrigation planning and management.

The limit to wheat water-use efficiency in eastern Australia. I. Gradients in the radiation environment and atmospheric demand

In the wheatbelt of eastern Australia, rainfall shifts from winter dominated in the south (South Australia, Victoria) to summer dominated in the north (northern New South Wales, southern Queensland). The seasonality of rainfall, together with frost risk, drives the choice of cultivar and sowing date, resulting in a flowering time between October in the south and August in the north. In eastern Australia, crops are therefore exposed to contrasting climatic conditions during the critical period around flowering, which may affect yield potential, and the efficiency in the use of water (WUE) and radiation (RUE). In this work we analysed empirical and simulated data, to identify key climatic drivers of potential water- and radiation-use efficiency, derive a simple climatic index of environmental potentiality, and provide an example of how a simple climatic index could be used to quantify the spatial and temporal variability in resource-use efficiency and potential yield in eastern Australia. Around anthesis, from Horsham to Emerald, median vapour pressure deficit (VPD) increased from 0.92 to 1.28 kPa, average temperature increased from 12.9 to 15.2°C, and the fraction of diffuse radiation (FDR) decreased from 0.61 to 0.41. These spatial gradients in climatic drivers accounted for significant gradients in modelled efficiencies: median transpiration WUE (WUEB/T) increased southwards at a rate of 2.6% per degree latitude and median RUE increased southwards at a rate of 1.1% per degree latitude. Modelled and empirical data confirmed previously established relationships between WUEB/T and VPD, and between RUE and photosynthetically active radiation (PAR) and FDR. Our analysis also revealed a non-causal inverse relationship between VPD and radiation-use efficiency, and a previously unnoticed causal positive relationship between FDR and water-use efficiency. Grain yield (range 1-7 t/ha) measured in field experiments across South Australia, New South Wales, and Queensland (n = 55) was unrelated to the photothermal quotient (Pq = PAR/T) around anthesis, but was significantly associated (r2 = 0.41, P < 0.0001) with newly developed climatic index: a normalised photothermal quotient (NPq = Pq . FDR/VPD). This highlights the importance of diffuse radiation and vapour pressure deficit as sources of variation in yield in eastern Australia. Specific experiments designed to uncouple VPD and FDR and more mechanistic crop models might be required to further disentangle the relationships between efficiencies and climate drivers.

Effect of timing of a deficit-irrigation allocation on corn evapotranspiration, yield, water use efficiency and dry mass

Water regulations have decreased irrigation water supplies in Nebraska and some other areas of the USA Great Plains. When available water is not enough to meet crop water requirements during the entire growing cycle, it becomes critical to know the proper irrigation timing that would maximize yields and profits. This study evaluated the effect of timing of a deficit-irrigation allocation (150 mm) on crop evapotranspiration (ETc), yield, water use efficiency (WUE = yield/ETc), irrigation water use efficiency (IWUE = yield/irrigation), and dry mass (DM) of corn (Zea mays L.) irrigated with subsurface drip irrigation in the semiarid climate of North Platte, NE. During 2005 and 2006, a total of sixteen irrigation treatments (eight each year) were evaluated, which received different percentages of the water allocation during July, August, and September. During both years, all treatments resulted in no crop stress during the vegetative period and stress during the reproductive stages, which affected ETc, DM, yield, WUE and IWUE. Among treatments, ETc varied by 7.2 and 18.8%; yield by 17 and 33%; WUE by 12 and 22%, and IWUE by 18 and 33% in 2005 and 2006, respectively. Yield and WUE both increased linearly with ETc and with ETc/ETp (ETp = seasonal ETc with no water stress), and WUE increased linearly with yield. The yield response factor (ky) averaged 1.50 over the two seasons. Irrigation timing affected the DM of the plant, grain, and cob, but not that of the stover. It also affected the percent of DM partitioned to the grain (harvest index), which increased linearly with ETc and averaged 56.2% over the two seasons, but did not affect the percent allocated to the cob or stover. Irrigation applied in July had the highest positive coefficient of determination (R2) with yield. This high positive correlation decreased considerably for irrigation applied in August, and became negative for irrigation applied in September. The best positive correlation between the soil water deficit factor (Ks) and yield occurred during weeks 12-14 from crop emergence, during the "milk" and "dough" growth stages. Yield was poorly correlated to stress during weeks 15 and 16, and the correlation became negative after week 17. Dividing the 150 mm allocation about evenly among July, August and September was a good strategy resulting in the highest yields in 2005, but not in 2006. Applying a larger proportion of the allocation in July was a good strategy during both years, and the opposite resulted when applying a large proportion of the allocation in September. The different results obtained between years indicate that flexible irrigation scheduling techniques should be adopted, rather than relying on fixed timing strategies.

Variation in ecophysiology and carbon economy of invasive and native woody vines of riparian zones in south-eastern Queensland

Exotic and invasive woody vines are major environmental weeds of riparian areas, rainforest communities and remnant natural vegetation in coastal eastern Australia, where they smother standing vegetation, including large trees, and cause canopy collapse. We investigated, through glasshouse resource manipulative experiments, the ecophysiological traits that might facilitate faster growth, better resource acquisition and/or utilization and thus dominance of four exotic and invasive vines of South East Queensland, Australia, compared with their native counterparts. Relative growth rate was not significantly different between the two groups but water use efficiency (WUE) was higher in the native species while the converse was observed for light use efficiency (quantum efficiency, AQE) and maximum photosynthesis on a mass basis (Amax mass). The invasive species, as a group, also exhibited higher respiration load, higher light compensation point and higher specific leaf area. There were stronger correlations of leaf traits and greater structural (but not physiological) plasticity in invasive species than in their native counterparts. The scaling coefficients of resource use efficiencies (WUE, AQE and respiration efficiency) as well as those of fitness (biomass accumulated) versus many of the performance traits examined did not differ between the two species-origin groups, but there were indications of significant shifts in elevation (intercept values) and shifts along common slopes in many of these relationships – signalling differences in carbon economy (revenue returned per unit energy invested) and/or resource usage. Using ordination and based on 14 ecophysiological attributes, a fair level of separation between the two groups was achieved (51.5% explanatory power), with AQE, light compensation point, respiration load, WUE, specific leaf area and leaf area ratio, in decreasing order, being the main drivers. This study suggests similarity in trait plasticity, especially for physiological traits, but there appear to be fundamental differences in carbon economy and resource conservation between native and invasive vine species.

Maximising profitability with limited water in cotton farming systems

Diminishing water supply, changing weather patterns and pressure to enhance environmental flows are making it imperative to optimise water use efficiency (WUE) on cotton/grain farming systems. Growers are looking for better strategies to make the best use of limited water, but it is still not clear how to best use the available water at farm and field scale. This research project investigated the impact of management strategies to deal with limited water supplies on the yield and quality of irrigated cotton and wheat. The objectives were: (1) to develop irrigation management guidelines for the main irrigated crops on the Darling Downs for full- and deficitirrigation scenarios, taking into account the critical factors that affect irrigation decisions at the local level, (2) to quantify the evapotranspiration (ET) of Bollgard II cotton and wheat and its relationship to yield and quality under full- and deficit-irrigation scenarios, and (3) to increase industry awareness and education of farming systems practises for optimised economic water use efficiency.Objective (1) was addressed by (A) collaborating with ASPRU to develop the APSFarm model within APSIM to be able to perform multi-paddock simulations. APSFarm was then tested by conducting a case study at a farm near Dalby, and (B) conducting semi-structured interviews with individual farmers and crop consultants on the Darling Downs to document the strategies they are using to deal with limited water. Objective (2) was addressed by (A) building and installing 12 large (1 m x 1m x 1.5 m) weighing lysimeters to measure crop evapotranspiration. The lysimeters were installed at the Agri-Science Queensland research station at Kingsthorpe in November 2008, (B) conducting field experiments to measure crop evapotranspiration and crop development under four irrigation treatments, including dryland, deficit-irrigation, and full irrigation. Field experiments were conducted with cotton in 2007-08 and 2008-09, and with wheat in 2008 and 2009, and (C) collaborating with USQ on a PhD thesis to quantify the impact of crop stress on crop evapotranspiration and canopy temperature. Glasshouse experiments were conducted with wheat in 2008 and with cotton in 2008-09. Objective (3) was addressed by (A) conducting a field day at Kingsthorpe in 2009, which was attended by 80 participants, (B) presenting information in conferences in Australia and overseas, (D) presenting information at farmers meeting, (E) making presentations to crop consultants, and (F) preparing extension publications.As part of this project we contributed to the development of APSfarm, which has been successfully applied to evaluate the feasibility of practices at the whole-farm scale. From growers and crop consultants interviews we learned that there is a great variety of strategies, at different scales, that they are using to deal with limited water situation. These strategies will be summarised in the "e;Limited Water Guidelines for the Darling Downs"e; that we are currently preparing. As a result of this project, we now have a state-of-the-art lysimeter research facility (23 large weighing lysimeters) to be able to conduct replicated experiments to investigate daily water use of a variety of crops under different irrigation regimes and under different environments. Under this project, a series of field and glasshouse experiments were conducted with cotton and wheat, investigating aspects like: (A) quantification of daily and seasonal crop water use under nonstressed and stressed conditions, (B) impact of row configuration on crop water use, (C) impact of water stress on yield, evapotranspiration, crop vegetative and reproductive development, soil water extraction pattern, yield and yield quality. The information obtained from this project is now being used to develop web-based tools to help growers make planning and day-to-day irrigation decisions.

Weed–pathogen interactions and elevated CO2: growth changes in favour of the biological control agent

In this study, we used Parthenium hysterophorus and one of its biological control agents, the winter rust (Puccinia abrupta var. partheniicola) as a model system to investigate how the weed may respond to infection under a climate change scenario involving an elevated atmospheric CO2 (550 μmol mol−1) concentration. Under such a scenario, P. hysterophorus plants grew significantly taller (52%) and produced more biomass (55%) than under the ambient atmospheric CO2 concentration (380 μmol mol−1). Following winter rust infection, biomass production was reduced by 17% under the ambient and by 30% under the elevated atmospheric CO2 concentration. The production of branches and leaf area was significantly increased by 62% and 120%, under the elevated as compared with ambient CO2 concentration, but unaffected by rust infection under either condition. The photosynthesis and water use efficiency (WUE) of P. hysterophorus plants were increased by 94% and 400%, under the elevated as compared with the ambient atmospheric CO2 concentration. However, in the rust-infected plants, the photosynthesis and WUE decreased by 18% and 28%, respectively, under the elevated CO2 and were unaffected by the ambient atmospheric CO2 concentration. The results suggest that although P. hysterophorus will benefit from a future climate involving an elevation of the atmospheric CO2 concentration, it is also likely that the winter rust will perform more effectively as a biological control agent under these same conditions.

Fingerprinting environmental conditions and related stress using stable isotopic composition of rice (Oryza sativa L.) grain organic matter

Sea level rise (SLR) is a primary factor responsible for inundation of low-lying coastal regions across the world, which in turn governs the agricultural productivity. In this study, rice (Oryza sativa L.) cultivated seasonally in the Kuttanad Wetland, a SLR prone region on the southwest coast of India, were analysed for oxygen, hydrogen and carbon isotopic ratios (delta O-18, delta H-2 and delta C-13) to distinguish the seasonal environmental conditions prevalent during rice cultivation. The region receives high rainfall during the wet season which promotes large supply of fresh water to the local water bodies via the rivers. In contrast, during the dry season reduced river discharge favours sea water incursion which adversely affects the rice cultivation. The water for rice cultivation is derived from regional water bodies that are characterised by seasonal salinity variation which co-varies with the delta O-18 and delta H-2 values. Rice cultivated during the wet and the dry season bears the isotopic imprints of this water. We explored the utility of a mechanistic model to quantify the contribution of two prominent factors, namely relative humidity and source water composition in governing the seasonal variation in oxygen isotopic composition of rice grain OM. delta C-13 values of rice grain OM were used to deduce the stress level by estimating the intrinsic water use efficiency (WUEi) of the crop during the two seasons. 1.3 times higher WUE, was exhibited by the same genotype during the dry season. The approach can be extended to other low lying coastal agro-ecosystems to infer the growth conditions of cultivated crops and can further be utilised for retrieving paleo-environmental information from well preserved archaeological plant remains. (c) 2015 Elsevier Ltd. All rights reserved.

增强UV-B辐射对暖温带落叶阔叶林下土庄绣线菊水分生理和氮素利用及形态特性的影响

在中国北方大部分地区，水分始终是影响植物生长和分布的最主要限制性因子之一，植物在其生长期经常遭受水分胁迫。不仅如此，随着大气同温层中臭氧浓度的减少，过量的有害紫外辐射（主要是UV-B，280nm-320nm）将穿透大气层达到地球表面。随着全球变化的加剧，这些地区的植物将不可避免地受UV-B和水分胁迫的共同作用。 本实验是在北京东灵山暖温带森林生态系统中，选择常见灌丛土庄绣线菊(Spiraea pubescens)，建立UV-B控制实验。连续三个生长季每天增补9.4kJ•m-2的辐射剂量，模拟臭氧衰减17%时近地表面UV-B辐射的增强。本实验的目的是观测在野外环境下，长时间人工增强UV-B辐射对土庄绣线菊水分生理、氮素利用以及形态特征的影响。具体对以下指标进行测定：叶片的气孔导度、碳同位素比率（δ13 C）、叶含水量、叶面积、水分利用效率（WUE）、叶全氮含量、叶氮素再吸收率。 实验结果表明，增强UV-B辐射显著减少了土庄绣线菊的叶面积（50.1%），提高了叶片全氮含量(102%)，处理植株的氮素再吸收率比对照植株高出50.9%。同时，UV-B辐射还在一定程度上（尽管统计显示不显著）降低了气孔导度(16.1%)、胞间CO2浓度与大气CO2浓度之比(Ci/Ca) (4.0%)、提高了碳同位素比率（δ13 C）(20.5‰)、叶含水量(3.1%)及比叶重（SLW）(5.2%)，从而导致水分利用效率（WUE）的增加(4.1%)，植物的抗旱能力增强。值得注意的是，深层土壤（30-40cm）含水量变化会影响气孔导度、δ13 C、WUE对紫外辐射的响应程度：在土壤干旱的季节（6月和9月），气孔导度、δ13 C、WUE这些指标处理和对照的差异很小，但是当土壤水分充足时（7月和8月），处理和对照的差异就较为显著。另外，随着实验处理时间的延长UV-B的效应变得不显著。相关分析表明，UV-B辐射降低了土壤含水量（30-40cm）与土庄绣线菊叶含水量、δ13 C、Ci/Ca、气孔导度的相关系数，增强了WUE与土壤含水量的相关性，这也许是由于UV-B辐射增强了WUE对土壤水分变化的敏感性。本研究的结果表明UV-B辐射对土庄绣线菊的形态和生长有显著的影响，但对主要水分生理指标影响不显著。

CO2倍增对紫花苜蓿的生态生理作用及模型

本论文是国家自然科学基金重大项目“中国陆地生态系统对全球变化的反应模式研究”下子项目“对全球变化反应植物生态生理学的基础模型研究”中的重要部分。 本文研究了紫花苜蓿(Medicago Sativa L．)在C02倍增下光合作用、蒸腾作用、气孔导度、叶面积、物候进程、高度、以及生物量的生态生理变化，并在此基础上对苜蓿进行了生态生理模型化的研究。 在倍增(694ppm)和对照(375ppm) C02浓度下，对紫花苜蓿的生态生理学的研究表明，以整个生育期计，倍增组的表观光合作用比对照组可提高18.7%：气孔导度略有下降(2%);蒸腾作用减少了2.7%;水分利用效率提高了30.1%;叶面积增加了48.9%;每株植物白天的净光合总量可提高76.7%，另外，植株高度和整株生物量的测定也显示了C02增加对苜蓿的正效应。 本文还对生理指标的实测数据进行了模型化的研究。对光合作用模型和气孔导度模型中参数的拟合结果表明，C02倍增下，苜蓿的光能转化效率(α)，电子传递速率(Jmax)比对照组都有明显的提高，最大气孔开度(Gsmax)略有下降．

西双版纳片段化热带雨林植物水分利用效率的边缘效应

本研究利用稳定性碳同位素法，测定了西双版纳城子片断化季雨林和补蚌沟谷雨林7条样线上70个科226种林下植物叶片δ13C值，并以此为表征研究了片断化雨林植物WUE边缘效应，结果表明： 西双版纳热带雨林林下植物叶片δ13C值与世界范围内其他热带雨林的研究结果相近。补蚌样地的植物叶片δ13C值显著低于城子样地植物叶片δ13C值，说明水分条件是植物水分利用效率的主要决定因素。 常绿植物的δ13C值显著低于落叶植物，由此可推知常绿植物的WUE显著低于落叶植物;乔灌草和藤本植物叶片δ13C值也存在显著差异，表现为：藤本＞灌木，乔木＞草本。 方位对边缘效应的影响不容视。东、西、南、北四个方位相比，边缘的产生对四面影响最大，北面最小。森林边缘对植物WUE的影响深度至少进入林内30m。 侵入物种所占比例在林缘较大，而进入林内锐减。有些植物δ13C值与距离显著负相关，说明这些植物对水分条件敏感（如假海桐，木奶果），若小气候条件继续变干，他们有在林缘消失的危险;有些相关不显著（如冬叶、锡金粗叶木、小叶藤黄），则植物对水分条件不敏感，这些物种在边缘产生后对环境条件变化影响不敏感，不会因森林片断化而迅速灭绝。

内蒙古锡林河流域主要植物种、功能群和群落水分利用效率的研究

近二十年来，碳同位素技术己被广泛应用于植物生态学，特别是植物“碳一水”关系的研究中。植物的碳同位素组成（δ13C值）是叶片组织合成过程中光合活动的整合，它反映了植物长期的水分利用效率。内蒙古锡林河流域位于我国温带典型草原的核心区域，水分是制约本区植物生产力和群落稳定性的限制因素。因此关于本区植物水分利用效率和水分利用状况的研究，对探讨植物对生境干旱化的适应与响应机制具有十分重要的理论和实践意义。本研究沿土壤水分梯度在锡林河流域选取了沼泽化草甸、盐化草甸、草甸草原、典型草原、退化草地和疏林沙地等8个代表性植物群落，研究主要植物种、功能群和群落的碳同位素组成及叶片含水量、脯氨酸含量等与植物抗旱性相关的生理指标的变化，从植物种、功能群和群落三个层次研究了不同水分条件下植物水分利用效率的变化及其对不同水分生境的响应与适应机制。 　　1）在所调查的8个植物群落中，C3植物占绝对优势;C3植物的δ13C值和水分利用效率越大，其在整个流域中的分布频度越高，生物量也越大;与生长在湿润生境中的植物相比，生长在较干旱生境中的植物能积累更高水平的脯氨酸。以上结果表明，锡林河流域的植物可能通过两种机制适应当地的干旱生境：一是通过调节气孔导度提高植物的水分利用效率;止是通过积累高水平的脯氨酸增强植株的渗透调节能力并维持相对稳定的水分含量。 　　2）依照生活型将锡林河流域主要植物种划分成6个植物功能群：乔木、灌木、半灌木、多年生禾草、多年生杂类草和一年生植物。在较湿润生境，多年生杂类草更加丰富并构成了群落地上生物量的绝大部分;而在较干旱生境下，多年生禾草在群落中起更重要的作用;随着土壤含水量下降，灌木和半灌木逐渐增多，且在退化草地和沙地中其相对生物量迅速增加;多年生禾草别3c值显著高于其它功能群;随着土壤水分可利用性降低，多年生禾草和杂类草的别3c值表现出增加的趋势，而灌木／半灌木则表现出相反的趋势。以上结果进一步证明了，在典型草原区以生活型为基础划分的植物功能群可以用来进行较大尺度植物一水分关系的研究。 　　3）依照植物的水分生态类群，将锡林河流域主要植物种划分为六个植物功能群：旱生植物、中旱生植物、旱中生植物、中生植物、湿中生植物和湿生植物。在较湿润生境中（沼泽化草甸和盐化草甸），湿中生和湿生植物成为优势种并构成地上生物量的主体;在干旱生境中（草甸草原、典型草原和退化草地），旱生和中早生植物占绝对优势并构成群落生物量的90％以上;随着不同水分生态类群所适应的生境从干旱到湿润逐渐转变，植物的δ13C值和水分利用效率显著降低;旱生植物叶片脯氨酸含耸最高，湿中生和湿生植物脯氨酸含量最低，不同水分生态类群脯氨酸含量与其δ13C值和地上生物星．显著正相关关系。 　　4）不同群落类型的平均δ13C值有显著不同，表现为：典型草原＞退化草地＞沙地＞退化恢复草地＞草甸草原之盐化草甸＞沼泽化草甸。C4植物的出现、不同物种δ13C值的差异和同一物种在不同生境下δ13C值的变化是影响群落平均δ13C值的主要因素，而这些因素与土壤水分状况和干扰历史（特别是放牧）密切相关。 　　此外，本文还研究了氮素添加对羊草和大针茅光合和水分利用效率的影响。土壤含氮量的增加可以显著提高羊草叶片光合能力和叭JE，而对大针茅的影响不大。作为锡林河流域两种优势植物，羊草和大针茅通过不同的生理机制来维持较高的WUE适应干旱生境：羊草为高光合、高蒸腾，而大针茅为低光合、低蒸腾。羊草较高的WUE是以降低氮利用效率 （NuE）为代价的;而大针茅在维持较高WUE的同时仍能维持较高的NUE，这一特征使大针茅可以广泛分布于更加干旱和贫瘩的地区。 　　以上研究结果，为深入开展典型草原生态系统植物与水分关系的研究提供了有价值的信息，进一步证实了稳定性碳同位素技术可以有效地指示不同群落类型中主要植物种长期水分利用效率。同时，通过对其它相关生理指标的测定，可以更好地探讨植物对水分限制的适应策略。我们的研究结果从植物种、功能群和群落三个层次进一步揭示了植物对干旱生境的适应机制，并初步阐明了人类干扰特别是过度放牧对草原群落建群种和优势种的生态替代或／和灌丛入侵的影响。这些研究对生物多样性保护、全球变化和区域可持续发展等热点问题的研究都具有重要的意义。在今后的研究中，结合其它稳定性同位素（如2H，18O和15N）技术，将有助于我们进一步深入研究蒙古高原植物对气候变化和过度放牧的适应与响应机制。 　　

半干旱区典型植物对气候变化与C02浓度升高的响应与适应

1.羊草对土壤水分的响应与适应 羊草生物量随着土壤水分含量的降低逐渐降低，后期的降低幅度远远大于前期。干旱促进鞘分配增加，增加了在处理初期的根的分配，但到后期则使之减少，表明羊草在经历较长期的持续干旱后通过增加根部的比重来提高抗旱性的能力逐渐降低。轻度(LD)、中度干旱(MD)对羊草叶片相对含水量(RWC)、气孔密度、光合参数、荧光猝灭参数和群体日交换速率无显著影响，但严重土壤干旱使它们显著降低。 羊草叶片的可溶性蛋白质以中度干旱的最高，严重干旱(SD)特别是极严重干旱(VD)使之显著降低，游离氨基酸含量(FAA)的变化与之相似。随着土壤水分含量的降低硝酸还原酶(NR)活性逐渐下降，而谷氨酰胺酶合成酶(GS)的活性变化则是LD和MD使之分别增加了25.75%和12.22%，SD和VD则分别减少了8.21%和28.72%，说明了NR的活性变化对土壤干旱较敏感，而GS的活性则对适度的干旱有一定程度的适应性。LD处理没有增加天冬酰胺酶(AE)和内肽酶(EP)两种酶的活性，但MD、SD和VD使两种水解酶的活性显著增加，说明轻度土壤干旱对蛋白质和氨基酸的分解作用有稍降低作用，但随着土壤干旱程度的加剧，又极大地促进了这个分解过程。严重和极严重土壤干旱显著降低了叶片的总核酸含量和RNA的含量，暗示严重程度的土壤水分胁追限制了核酸的合成代谢，加强了其分解代谢，严重土壤干旱还显著增加了丙二醛(MDA)的含量，说明提高了羊草叶片叶肉细胞的膜质过氧化水平。 2.羊草对土壤干旱和复水的响应与适应 羊草受到适当的干旱驯化可促进生长，但过长时间的干旱处理，复水后未能补偿损失的生物量和叶面积。羊草叶片的气孔密度以中度干旱持续期(Mtd)处理的最高，其次是短期干旱持续期( Std)，二者分别比没有经过土壤干旱的处理（对照）增加了14.90%和3.61%，但长期干旱持续期(Ltd)却使之减少了27.19%，气孔指数亦有类似的趋势。复水增加羊草叶片的光合速率、气孔导度、蒸腾速率，近期复水的激发效应明显大于前期，而对夜晚的呼吸作用影响不显著。水分利用率( WUE)的日变化动态呈“M”字型曲线，以Mtd的WUE值的峰值最大，以三次曲线拟合WUE的24小时日进程最佳。叶绿素荧光动力学的分析结果表明，复水，特别是最近的复水可显著改善羊草叶片的PS II性能，增加叶绿素a，b的含量及其比值，提高碳酸酐酶的活性。 羊草含氮量以叶片的最高(4.40%)，根部的最低(1.99%)，枯叶、茎鞘和根茎的含量差异较小(2.26～4.40%)。所有器官的含氨量对土壤水分处理的响应基本一致，以对照处理的最低，Std的最高。各器官的碳氮比都是以对照的最高，而其它土壤水分处理相差不显著，给于一定时间的土壤干旱处理可使羊草获得较强的氨代谢能力。Std的氮素总拥有量最多，和对照相比，绿叶、枯叶、茎鞘、根茎和根分别提高了35.58、26.88、23.49、31.66、40.75%，而Ltd的含氨总量呈下降趋势，说明短时间的土壤水分干旱处理可明显促进羊草各器官和植株的氮素积累，而较长时间的土壤干旱则不利于氮素的积累。羊草各器官氮素绝对量占整株的百分比从大到小依次为：绿叶(42.42-44.00%)、根茎(20.13—23.69%)、根(15.43～17.18%)、枯叶(10.07～11.30%)和茎鞘(7.27～8.67%)，表明叶片的氮素存量占植株的一半以上。Mtd处理增加了叶片的氮素贡献率，减少了茎鞘和根茎的贡献率，有利于加强叶片的光合性能。 以中度干旱持续期(Mtd)处理的叶片可溶性蛋白质含量、谷氨酰胺合成酶(GS)活性、RNA含量为最高，但中长期的土壤干旱处理再复水后则显著降低了羊草叶片内肽酶(EP)的活性和MDA的含量，说明给于一定时间的土壤干旱处理可使羊草叶片保持较高的蛋白质代谢水平，降低膜脂过氧化水平。 3.羊草对昼夜温差与土壤水分交互作用响应与适应 昼夜温差减少使单株羊草的生物量降低21.3%，分蘖和根的生物量减少，而鞘的生物量稍增加，显著降低了严重和极严重条件下的生物量。温差缩小降低了分蘖和根的投资比例，减少植株的地下部分生物量，而增加新叶、鞘和分蘖光合产物的比例，表明温差的减少将抑制光台产物向地下部分的转移。温差减少对充足土壤水分和轻度干旱处理的放射性比强影响较小，但减少了其它3种干旱处理的放射性比强。其原因主要是减少了植株鞘、根和根茎的放射性比强，显著增加了饲喂叶和心叶的放射性比强，表明温差的缩小阻止了“源”的光合产物向“库”的转移，降低对分蘖和根的投资，不利于羊草对干旱逆境的适应。 昼夜温差缩小使羊草叶片的气孔密度降低4.01%，而且减少了土壤干旱对气孔密度的影响。较高的昼夜温差和较低的昼夜温差相比，羊草叶片的光合速率和WUE分别增加了7.37%和20.09%;而气孔导度、胞间CO2浓度和蒸腾速率分别降低了14.03%、2.57%和10.80%。昼夜温差减少降低了土壤干旱的δ13C值，说明可能减少处在干旱条件下的植株WUE，暗示减小昼夜温差不利于增大羊草叶片对土壤水分亏缺的耐性。昼夜温差的缩小主要影响了下午羊草群体的CO2交换速率，增加了对照、LD和MD处理的夜间呼吸速率，降低了尤其是显著降低了在土壤缺水条件下的CO2昼夜净交换量，而在两个昼夜温差条件下都是以LD的最高。 昼夜温差缩小影响氮素含量在羊草各器官中的氮素分配，降低了叶片和茎鞘中的氮素百分比含量，但增加了根茎和根中的氮素含量。在较小的昼夜温差条件下，LD显著增加了叶片的氮素含量，但其它土壤水分处理影响不显著，在较大的昼夜温差条件下，亦是LD显著增加了叶片的氮素含量，但MD也使其显著增加。昼夜温差缩小增加了叶片和茎鞘的碳氮比，降低了根茎和根的碳氮比，使叶片在中度和严重土壤水分胁迫时无变化，降低了响应于土壤水分变化的调节弹性。 昼夜温差缩小使羊草叶片可溶性蛋白质降低了16.14%。LD和MD显著促进叶片可溶性蛋白质含量增加，SD和VD都显著地降低了羊草叶片的可溶性蛋白质含量。昼夜温差缩小有使羊草叶片游离氨基酸降低的趋势，主要是降低土壤干旱条件下的游离氨基酸含量，虽然昼夜温差缩小稍增加了可溶性耱的含量(4.68%)，但使土壤干旱激发效应变得不显著，表明温差的缩小降低了叶片中渗透调节物质的积累，不利于羊草抗旱性的提高。昼夜温差缩小显著降低了羊草叶片的NR、GS和谷氨酸脱氢酶(GDH)的活性，在较小昼夜温差条件下几乎看不出土壤水分降低的激发效应，而在高温差下则明显看出LD的激发效应。昼夜温差减少加强土壤干旱对天冬酰胺酶和内肽酶活性的影响，加强了蛋白质和关键氨基酸的分解作用，提高了叶片的膜脂过氧化水平，不利于羊草对干旱的适应性响应。 在土壤水分充足条件下，昼夜温差缩减对羊草叶片气孔形态的影响不明显，但可以看出温差较大时有较多的星状蜡质覆于气孔表面及其周围，这可能是由于夜间温度升高后植物为了减少水分的散失而采取的一种适应性策略。在本实验条件下，中度和严重土壤水分胁迫使气孔变得更加凹陷，气孔更加坚挺，体现了对土壤干旱的适应性反应。对羊草叶片叶肉细胞超微结构的观察表明，昼夜温差对土壤水分充足时的超微结构影响不显著，但在严重土壤干旱条件下，昼夜温差缩减似乎减少了叶绿体中的淀粉粒，加速了叶绿体和线粒体膜的裂解，表明昼夜温差缩减加大了严重土壤干旱对羊草叶片叶肉细胞超微结构的负面影响。 4.羊草对温度和土壤水分交互作用的响应与适应 温度升高使羊草叶片气孔导度和蒸腾速率增加，使光合速率(A)和水分利用率( WUE)降低。土壤干旱和高温均导致最大光化学效率、量子产额和光化学荧光猝灭系数降低，使非光化学荧光猝灭系数升高。土壤干旱减少了羊草幼苗的生物量，却显著增加了根的贡献率和根冠比，而高温使二者显著地降低。在本实验条件下，直到极端干旱才显著降低了羊草叶片的A和WUE，而其他的水分处理影响不显著。土壤干旱使得叶片中的含氮量显著降低：温度对叶片氮含量无显著影响，但却显著地降低了根中的氮含量，尤其显著增加根与叶片氮含量的比率。高温加强了干旱对光合性能的影响，表明高温降低了羊草对干旱的适应能力。 温度过低或过高都对叶片保持高水平的可溶性蛋白质不利，温度过高还削弱了土壤水分亏缺对叶片中游离氨基酸的激发作用。水分梯度对20℃时的GS活性无显著影响，但LD促使26℃和29'C下的GS活性增加，MD以上强度的土壤干旱都显著降低了23℃以上温度尤其是29℃和32℃时的GS活性，表明高温和干旱的协同效应对谷氨酰胺的合成不利。高温增加了AE和EP的活性，加强了土壤干旱对羊草叶片蛋白质和氨基酸分解的促进作用，在一定的土壤水分条件下，高温对RNA的合成作用增强，认为这是对高温胁迫的一种适应性响应。 随着温度的升高羊草叶片中的可溶性糖含量逐渐增加，土壤干旱亦增加了其可溶性糖的含量，但至极端干旱时则使之降低。在20℃下，土壤水分对羊草叶片的可溶性糖的含量无显著影响，在23—32℃温度条件下则是土壤干旱增加了其值，但至VD时则使之显著降低。不同温度条件下，土壤水分对羊草叶片的MDA影响不同，在20℃下，无显著影响，在23和26℃下，SD和VD使MDA稍升高，但在29和32℃条件下使之显著增加，说明温度升高加强了土壤干旱所引发的增加叶片膜质过氧化水平的负面影响。 5.柠条和杨柴对CO2浓度倍增和土壤水分交互作用的响应与适应 土壤干旱使拧条和杨柴的生物量在倍增CO2浓度条件下比在正常浓度条件下降低幅度更大。CO2浓度倍增较大地促进了充足水分条件下的植物生长，而对干旱条件下的生长促进作用则较小。无论在中度条件下还是在严重干旱条件下，两种优势植物均是在倍增CO2浓度条件下增加的根冠比幅度较大;无论在中度条件下还是在严重干旱条件下，且无论正常CO2浓度条件下还是在倍增CO2浓度条件下均是杨柴增加的幅度大。CO2浓度倍增主要增加了水分充足和MD的单位叶面积质量(LMA)，但反而降低了严重干旱的LMA。 CO2倍增使δ13C降低，但土壤干旱使之增加。用“库”（根）中的δ13C值对“源”（叶片）中的δ13C作图，可以用以评价碳分配以及“库”中新增生物量，两种沙生灌木叶片与根部的δ13C值呈极显著线性关系，杨柴的斜率大于柠条的，表明前者叶片与根部在光合产物分配上具有较高的可塑性，这和干旱条件下杨柴的根冠比增加相关联。杨柴的“源库”调节特性反映了对逆境具有较高的耐性。 CO2倍增使柠条和杨柴叶片含氮量分别降低了10.40%和5.06%，土壤干旱有使柠条叶片含氮量增加的趋势，但中度干旱没有增加羊柴叶片的含氮量。CO2倍增使叶片的碳氮比显著增加，而干旱使之降低。CO2浓度倍增降低叶肉细胞质膜的过氧化产物MDA的含量，干旱亦使叶片的MDA含量增加。叶片含氮量与MDA呈显著正相关，表明CO2倍增有保护叶片免受土壤干旱的作用，但干旱的负面影响是CO2倍增效应所难以弥补的。 CO2倍增降低了柠条叶片的可溶性蛋白质的含量，但在干旱条件下降低幅度较小，说明CO2浓度升高条件下可减轻干旱影响叶片中可溶性蛋白质的强度，体现了CO2浓度倍增对植物的抗旱性有利的一面。CO2浓度倍增使土壤水分充足条件下的柠条叶片中游离氨基酸含量降低17.24%，却使SD条件下增加10.78%，表明土壤干旱导致的叶片游离氨基酸含量的增加平衡了CO2升高造成的降低。在充足土壤水分和MD条件下，CO2浓度倍增对核酸总含量和RNA含量有稀释效应，但严重干旱条件下，CO2倍增提高了核酸总含量和RNA的含量。

夏玉米不同生育期对干旱胁迫的响应与适应机制

干旱化问题将在全球环境变化下进一步加剧，并可能严重影响玉米。玉米是我国主要粮食和最重要的饲料作物，其重要性日益突出。水分是制约玉米产量的关键因子。为此，本研究利用大型活动遮雨棚对玉米进行了出苗后全程水分控制试验，研究大田条件下玉米不同生育期对不同土壤水分（包括水分充足well-watered, WW;适度干旱moderately stressed,MS;和严重干旱severely stressed,SS）的响应及适应机制。研究结果表明： 在吐丝和籽粒形成期，Ms对叶片相对含水量和相对电导率的影响没有达到显著或极显著水平，而SS则极其显著地降低叶片相对含水量和增加质膜透性。并且干旱胁迫下，夏玉米生育进程中保护酶SOD、POD和CAT活性基本呈现一致下降的态势，膜脂过氧化作用增强。短期干旱胁迫对SOD）和POD（在第十三叶期）保护酶有一定的激发效应，但此效应维持不长，其后骤降。 干旱会引起叶绿素a，b含量及总叶绿素含量的减少。MS下营养阶段的叶绿素含量没有明显变化，但随着MS的延续，叶绿素含量在生殖阶段显著降低。而SS的叶绿素含量最初就呈现降低并逐渐扩大。另外，在干旱胁迫下叶净光合速率(PN)和蒸腾速率(E)的降低因干旱强度和时间以及发育阶段而异，而且Ss所引起的不利影响更为凸现。SS显著降低营养和生殖阶段的水分利用效率( WUE)，然而MS基本导致前中期WUE增加，后期则减少。 土壤干旱胁迫下，绿色LAI明显降低，特别是生殖时期最高穗位叶面积显著降低：地上部生物量积累在各生育期均为减少。而且，干旱显著减少各生育期的根干重，但MS对第十八叶期(V18)的根干重有短期的促进作用。干旱胁迫下，根冠比在不同生育时期有增有减。MS对第十七叶期(V17)的叶面积、抽雄吐丝出现、叶片展开、最终叶片数以及收获指数影响不大，但其却显著减少各阶段株高、叶面积（第十七叶期除外）、茎粗和生物量积累。随着MS的延续，产量性状诸如穗粒数、百粒重均为降低，而SS对各生育阶段所有生长特性、产量性状及收获指数的影响都较MS更为不利。 生育前期遭遇干旱，可使叶片展开明显迟缓，并且最终叶片数减少。尤其SS减少最终叶片数1～2片，并且延迟抽雄4—5 d，吐丝4—5 d，从而可能导致成熟期推迟。 植物器官的营养吸收动态在短期干旱作用和长期作用之间有所不同。而且P和K元素的积累方式也有别。基本上，干旱胁迫显著降低植物器官在不同生育期的全P和K元素的吸收，尽管后期一些器官诸如叶、鞘和茎等的吸收有所增加，特别是干旱严重影响了根的吸收能力，而且SS较MS对全P.K吸收影响更甚。总之，干旱所导致的生物量减少与植物器官的全P、K吸收的减少是相伴而生的。 与WW相比较，MS和SS的产量两年内分别降低了20,4%—26,1%和59.2%～84.5%，穗粒数分别降低了12.1%～19.7%和39,8%～88.1%，以及百粒重分别降低了2.1%~2.7%和17.7%—46,9%。研究进一步表明，干旱胁迫对多数玉米籽粒的营养品质有利。与WW相较而言，N含量、可溶性总糖、可溶性还原糖、Zn. Ca. Cu. Mg和Mn元素在MS下分别提高了5.9%，39-0%，97.5%，12.1%，4.4%，7.5%，6.1%和2.9%，而在SS下则分别提高了8.6%，99.3%，300.0%，27.8%，24.0%，1 5.3%，9.8%和7.9%。但是，一些玉米籽粒的营养品质诸如淀粉、P和K含量却受到干旱胁迫的不利影响，与WW相比较，MS使籽粒淀粉、P和K含量分别降低了8.3%．12.6%和3.7%，而SS则分别降低了33.3%, 14.6%和18.6%。粗脂肪含量则表现有所不同，与WW比较而言，MS对之有利，2年平均增加9.2%，而SS对之不利，2年平均减少11.3%。 总之，玉米生理生态特征、地上部各部分干物质生产、根系生长、营养吸收、产量性状、营养品质对干旱胁迫的响应和适应不仅依赖于干旱的严重程度（包括强度和时间），而且也依赖于玉米发育阶段。本研究认为，在半湿润地区水分缺乏的条件下，有限灌溉（最低土壤相对含水量55%士5%）在营养阶段抽雄前实施可行。