3种荒漠灌木的用水策略及相关的叶片生理表现

以新疆古尔班通古特沙漠南缘原始盐生旱生荒漠的3种建群灌木多枝柽柳(Ｔamarix ramosissima)、梭梭(Ｈaloxylon ammodendron)和琵琶柴(Reaumuria soongorica)为对象,跟踪自然降雨过程,利用LI-6400光合作用系统和Model 3005植物水分压力审测定光合作用和叶水势的变化,以研究浅层土壤水分条件改变对荒漠灌木主要叶片生理特征的影响;并在原始生境中将植株根系完整地深挖取出,进行根系形态结构调查,以确定此3种灌木根系功能型与用水策略。当浅层土壤分别处在水分充足及匮缺的条件下时．测定3种灌木的光合作用响应曲线和日过程曲线．以及黎明前和止午叶水势,结果表明：浅层土壤水分状况变化时,3种灌木的光合能力均没有显著改变;多枝柽柳的叶水势亦没有明显波动;而梭梭和琵琶柴的叶水势却表现出显著差异。在两种功能型根中,多枝柽柳为深根型,生存和乍理活动的维持主要依赖于地下水;而梭梭和琵琶柴为非深根型植物,主要水源是降水形成的浅层土壤水,其用水策略是根据水分条件行效调节根系和冠层生长,从而维持正常的光合作用。即荒漠灌木在长期适应的过样中．已形成不同的根系功能型和用水策略;叶水势对浅层土壤水分状况变化的种间差异性响应在一定程度上反映了这一点。同时．此3种荒漠灌小通过不同的个体适应策略都能够实现水分平衡和碳收支的有效调节,这主要体现为浅层土壤水分条件变化时光合响应的种间一致性。     

共生条件下三种荒漠灌木的根系分布特征及其对降水的响应

以全根系挖掘法,对共生于原始盐生荒漠生境中的多枝柽柳[Tamarix ramosissima (Ledeb.)]、梭梭[Haloxylon ammodendron(C. A. Mey.)Bunge]、琵琶柴[Reaumuria soongorica (Pall.) Maxim.]的根系分布特征进行了研究;对降水引发的湿润-干旱周期中植物同化枝水势、蒸腾速率的变化过程进行了跟踪观测,并据此计算3种植物的水分胁迫效应指数和土壤-植物系统导水度,以最终确定3种植物用水策略和其对降水的响应特征.研究结果表明,多枝柽柳的吸收根系分布范围从地下50cm到310cm,单株平均总吸收根表面积为30249.2cm2;梭梭的根系分布范围0～250cm,单株平均总吸收根表面积12847.3 cm2;琵琶柴的根系分布范围0～80cm,单株平均总吸收根表面积361.8 cm2.多枝柽柳为深根植物,主要利用地下水和深层土壤水,在降水引发的湿润-干旱周期中,其植物水分生理参数对降水无响应.琵琶柴为浅根植物,对降水响应极为显著.梭梭的根系分布特征介于多枝柽柳和琵琶柴之间,对地下水和降水都有利用,对降水响应显著.3种荒漠灌木对降水的响应差异显然与其根系分布、水分利用策略密切相关,在未来降水发生变化的情景下,根系分布特征的差异将决定着植物在水分资源竞争中的地位.具有较强根系形态可塑性的物种,如梭梭,将具有明显的竞争优势.     

准噶尔盆地东南缘多枝柽柳、白刺和红砂水分来源的异同

荒漠生态系统中, 水是植物生长最主要的限制因子。为了比较同一生境下不同荒漠植物的水分来源特征, 选取了同一生境下的多枝柽柳(Tamarix ramosissima)、白刺(Nitraria sibirica)和红砂(Reaumuria soongorica), 测定了这3种植物茎水和各潜在水源(降水、土壤水和地下水)的氢、氧稳定同位素比率(δD和δ¹⁸O)值, 并利用IsoSource软件计算了3种植物对潜在水源的利用比例。结果表明: 红砂和白刺的茎水δD和δ¹⁸O值及其水分来源有明显的季节波动特征。其中, 红砂为浅根系植物, 春季(3-5月)以表层土壤水为主要水源, 夏秋季节(6-10月)表层土壤含水量显著降低, 其主要的水分来源逐渐偏向于较深层的土壤水; 白刺的根系分布范围介于红砂和多枝柽柳之间, 在春季能够较多地利用表层土壤水, 而到了夏秋季节, 所利用的水分更多地来源于深层土壤水或地下水; 多枝柽柳为深根系植物, 其90%以上的水分来源于深层土壤水和地下水, 而且茎水δD和δ¹⁸O值及其水分来源没有季节波动特征。3种植物水分来源特征的差异与其水分利用策略密切相关, 同时, 也说明荒漠灌木可以通过自身调节向着最优(最有利)表现型发展, 从而最大限度地获取水分。     

干旱区水力提升的生态作用

水力提升是某些植物通过深层根系从较湿的深层土壤中吸取水分再通过浅层根系在较干的浅层土壤中释出的过程。水力提升所释出水量及其释出过程具有积极的生态学意义,它不但为伴生植物提供新的水源、改善其邻近植物的蒸腾作用、缓解水分亏缺、提高水分利用效率．而且促进植物的养分吸收、促进上层干土中分解、硝化和矿化等生化过程．尤其是发生在克隆植物植株间的水分共享这种特殊的水力提升,还影响群落的形成。然而,对于一般的水力提升而言,水力提升所释出水量到底有多少。其生态学作用有多大．有待深入研究。结合研究实践认为．要研究水力提升具有多大的生态学作用．必须准确地区分水;勺提升的水量和毛管水的上升水量。还建议了研究水力提升新的研究方法。     

土壤深层供水对冬小麦干物质生产的影响

采用根系研究装置研究了土壤深层供水对冬小麦干物质生产的影响 .结果表明 ,上层低湿度下层高湿度的处理在小麦灌浆期仍然保持了较高的土壤和叶片含水量 ,具有发达的根系 ,特别是 1m以下的根量在 4个处理中为最高 ,旗叶和穗的干重也最大 ,具有最大的产量潜力 .本研究表明 ,上层土壤较干下层土壤湿润有利于发挥小麦根信号的积极作用 ,平衡水分利用 ,同时通过对土壤水分的合理调节可以促进深层根的发育 ,有利于提高产量和水分利用效率 .     

植物根系提水作用研究述评

根系提水作用是植物在干旱生境下,通过根系将深层湿润土壤中的水分提升至浅层干燥土壤中的一种生理过程,不同植物具有不同的根系提水强度。这一研究的基本方式是分根法,而时域反射仪(TDR)法和中子水分仪法则是近年来较受重视的测定方法,并逐步由室内测定向大田测定过渡。研究的主要内容可归纳为五个方面,其中在根系提水作用与植物蒸腾作用、浅层土壤营养的有效性以及与植物遗传性的关系等方面,研究结果较为一致,但在提水的数量和发生提水作用时土壤水分状况等方面,研究结果不尽相同;对植物光合、呼吸等其他代谢的影响以及室内测定结果与大田条件一致性等问题的研究,报道较少,特别是作用机理的研究。因此,深入这方面的研究对于进一步提高植物水分利用率,促进旱地节水农业和畜牧业的发展具有重要的意义。     

新疆古尔班通古特沙漠南缘多枝柽柳光合作用及水分利用的生态适应性

光合作用和水分利用是陆地生态系统中限制植物光合同化和个体生长发育的重要因素。为了从根本上认识到荒漠绿洲过渡带植物对特殊生境和气候变化的响应和适应机制,有必要深入的了解植被对不同环境因子适应过程的协调性。本文以新疆古尔班通古特沙漠南缘荒漠绿洲过渡带的建群种多枝柽柳（Tamarix ramosissima）为研究对象,分别在自然状态,模拟降水、遮光处理以及降水和遮光双因子处理下,利用LI-6400光合作用系统和Model 3500植物水分压力室分别测定光合作用和枝条水势,研究多枝柽柳枝条水势对浅层土壤水分和光照变化的响应,以及在不同条件下的水分利用效率（WUE）和光能利用效率（LUE）的日进程。结果表明：这四种处理方式下,枝条水势随土壤水分的变化没有明显的差异,趋势表现为黎明前枝条水势最高（-1.2 MPa）,正午枝条水势最低（-3.2 MPa）,太阳落山后枝条水势逐渐升高的趋势;WUE和LUE日变化呈现出近似双峰型抛物线,WUE日变化曲线在11:00和16:00有两个不明显的波峰,LUE日变化曲线在10:00和16:00出现两个明显波峰。当光照发生变化时,遮光处理下的WUE和LUE日变化都较自然状态下的日变化低,分别降低了1.5 ?mol CO2?mmol-1 H2O的WUE和0.20*10-2 ?mol CO2??mol-1APAR的LUE。研究表明,作为深根系植物的多枝柽柳,生存主要依靠地下水来补给水分,所以浅层土壤水分条件变化没有明显引起光合和蒸腾的响应。而午间光强、高温是构成多枝柽柳光抑制现象的主要原因,这是多枝柽柳长期在荒漠绿洲过渡带形成的对抗环境胁迫的一种表现。     

光合有效辐射与地下水位变化对柽柳属荒漠灌木群落碳平衡的影响

全球气候变化和人类活动的加剧, 正导致古尔班通古特沙漠南缘原始盐生旱生荒漠地区的地下水位发生显著改变。大气污染导致该地区太阳辐射减少。以盐生荒漠建群种多枝柽柳(Tamarix ramosissima)为研究对象, 选择地下水位在2.9-4.5 m波动的典型原始生境, 观测了生长期内光合有效辐射和地下水位变化时的光合作用、蒸腾作用和叶水势等生理活动的季节变化, 调查了根系分布特征; 并利用涡度相关系统测定了生态系统碳水通量, 估算群落碳同化能力、蒸腾耗水量与叶面积指数的季节变化, 旨在揭示光合有效辐射和地下水位等环境因素对柽柳属(Tamarix)荒漠灌木群落光合作用的影响。研究结果表明: 降水造成的潜土层水分状况变化对多枝柽柳的碳平衡没有显著影响。深根系与气孔调节是多枝柽柳碳平衡适应荒漠环境水分状况的两个关键机制。特殊的气孔行为体现了多枝柽柳以高水分消耗为代价将其碳获取最大化的适应对策; 多枝柽柳生理与群落尺度的水分平衡和碳获取均依赖于深根系获取的稳定地下水源, 缓和的地下水位波动将不会扰动其现有的碳/水平衡, 地下水位剧烈下降将危及多枝柽柳的生存。此外, 光合有效辐射是另一个主要影响因素, 与群落碳获取呈显著正相关关系。群落碳同化能力的季节变化是光合有效辐射和地下水位共同影响下光合作用物候学特征的体现。过度开采地下水和直接破坏原生植被的行为, 将会严重地干扰多枝柽柳群落的生存, 进而破坏该区域现有的生态水文过程。     

刺槐根系对深层土壤水分的影响

对黄土高原主要造林树种刺槐根系及其林地土壤水分进行调查和监测,结果表明,在阴坡立地上,刺槐细根在距树干2.0 m范围内的水平分布无明显差异,最大分布深度均为2.0 m;而阳坡立地上刺槐细根在距树干0.5 m处的垂直分布深度可达2.0 m,且阴坡立地上细根密度特征值明显大于阳坡.根系对土壤深层水分的影响范围因不同立地条件下根系分布空间差异而不同,在阳坡立地上,刺槐根系对深层土壤水分的影响深度可达2.7 m处的土层,而在阴坡立地上,这种影响范围可达3.3 m处的土层.     

土壤水分和氮磷营养对冬小麦根系生长及水分利用的调节

模拟试验研究结果表明：在土壤相对含水量为４０％￣７０％范围内,水分亏缺严重,根水势和蒸腾蒸发量显著降低,根系生长严重受阻,根长变短,根干重降低,随着土壤水分趋于良好,根水势和蒸腾蒸发量显著增加,根干重在土壤相对含水量为５５％￣６２％之间时最大,而土壤相对含水量在５５％上下时根长达最长;土壤水分趋于轻度干旱有利根系下扎,土壤水分趋于良好利于根量增长。氮磷营养对小麦根系生长具有明显的调节作用。磷营养可     

Below-ground competition between trees and grasses may overwhelm the facilitative effects of hydraulic lift

Under large East African Acacia trees, which were known to show hydraulic lift, we experimentally tested whether tree roots facilitate grass production or compete with grasses for below‐ground resources. Prevention of tree–grass interactions through root trenching led to increased soil water content indicating that trees took up more water from the topsoil than they exuded via hydraulic lift. Biomass was higher in trenched plots compared to controls probably because of reduced competition for water. Stable isotope analyses of plant and source water showed that grasses which competed with trees used a greater proportion of deep water compared with grasses in trenched plots. Grasses therefore used hydraulically lifted water provided by trees, or took up deep soil water directly by growing deeper roots when competition with trees occurred. We conclude that any facilitative effect of hydraulic lift for neighbouring species may easily be overwhelmed by water competition in (semi‐) arid regions.     

Hydraulic lift: consequences of water efflux from the roots of plants

Hydraulic lift is the passive movement of water from roots into soil layers with lower water potential, while other parts of the root system in moister soil layers, usually at depth, are absorbing water. Here, we review the brief history of laboratory and field evidence supporting this phenomenon and discuss some of the consequences of this below-ground behavior for the ecology of plants. Hydraulic lift has been shown in a relatively small number of species (27 species of herbs, grasses, shrubs, and trees), but there is no fundamental reason why it should not be more common as long as active root systems are spanning a gradient in soil water potential (Ψ_s) and that the resistance to water loss from roots is low. While the majority of documented cases of hydraulic lift in the field are for semiarid and arid land species inhabiting desert and steppe environments, recent studies indicate that hydraulic lift is not restricted to these species or regions. Large quantities of water, amounting to an appreciable fraction of daily transpiration, are lifted at night. This temporary partial rehydration of upper soil layers provides a source of water, along with soil moisture deeper in the profile, for transpiration the following day and, under conditions of high atmospheric demand, can substantially facilitate water movement through the soil-plant-atmosphere system. Release of water into the upper soil layers has been shown to afford the opportunity for neighboring plants to utilize this source of water. Also, because soils tend to dry from the surface downward and nutrients are usually most plentiful in the upper soil layers, lifted water may provide moisture that facilitates favorable biogeochemical conditions for enhancing mineral nutrient availability, microbial processes, and the acquisition of nutrients by roots. Hydraulic lift may also prolong or enhance fine-root activity by keeping them hydrated. Such indirect benefits of hydraulic lift may have been the primary selective force in the evolution of this process. Alternatively, hydraulic lift may simply be the consequence of roots not possessing true rectifying properties (i.e., roots are leaky to water). Finally, the direction of water movement may also be downward or horizontal if the prevailing Ψ_s gradient so dictates, i.e., inverse, or lateral, hydraulic lift. Such downward movement through the root system may allow growth of roots in otherwise dry soil at depth, permitting the establishment of many phreatophytic species. Received: 2 June 1997 / Accepted: 24 September 1997     

胡杨根系水力提升作用的证据及其生态学意义

生长在塔里木河流域的荒漠河岸林植被虽长期忍受着高温和干旱的威胁, 然而它们却能够一直延续并保存至今。除了植物深根系吸水作用外, 另一个更主要的原因可能就是荒漠河岸林植被存在水力提升的效应。该文采用HRM热比率法茎流仪对3株胡杨(Populus euphratica)主根和侧根的液流速率分别进行了为期4 d的连续监测; 利用自动气象站对微气象因子：风速、空气相对湿度、叶面温度和地表温度进行连续监测; 同时采用了烘干法对不同深度土层在不同时刻的土壤含水率进行了取样分析。试验结果表明：胡杨主根液流在白天和夜间均表现为正值, 相反的, 胡杨侧根液流速率则出现了明显的夜间负向流动。胡杨根系0~120 cm土层土壤水分含量具有下湿上干的变化趋势; 胡杨侧根在夜间发生负向流动后, 土壤含水率显著升高, 尤其在60~120 cm土层中, 4:00土壤含水率上升幅度达到4:00时刻土壤含水率的22%~26%。影响胡杨侧根液流速率的主要气象因子主要是叶面水汽压亏缺。     

Soil strength and water content influences on corn root distribution in a sandy soil

Initial field observations revealed a shallow corn (Zea mays L.) root system on a Zimmerman fine sand in a corn/soybean (Glycine max L.) rotation. Since root distribution influences crop water and nutrient absorption, it is essential to identify factors limiting root growth. The objective of this study was to determine the factor(s) limiting corn rooting depth on an irrigated fine sand soil. Bulk density, saturated hydraulic conductivity, and soil water retention were measured on undisturbed soil cores. Corn root distribution assessed at tasseling over a 3-yr period showed an average of 94% of total root length within the upper 0.60 m of soil with 85% in the upper 0.30 m of soil. Mechanical impedance was estimated with a cone penetrometer on two dates with differing water contents. Cone penetrometer measurements greater than 3 MPa indicated mechanical impedance in soil layers extending from 0.15 to 0.35 m deep. Penetration resistance decreased as soil water content increased. However, soil water contents greater than field capacity were required to decrease penetration resistance below the 3 MPa threshold. Such water saturated conditions only occurred for short periods immediately after precipitation or irrigation events, thus roots usually encountered restrictive soil strengths. The soil layer from 0.15 to 0.60 m had high bulk density, 1.57 Mg m^-3. This compacted soil layer, with slower saturated hydraulic conductivities (121 to 138 mm hr-1), held more water than the soil above or below it and reduced water movement through the soil profile. Crop water use occurred to a depth of approximately 0.75 m. In conclusion, a compacted soil layer confined roots almost entirely to the top 0.60 m of soil because it had high soil strength and bulk density. The compacted layer, in turn, retained more water for crop use.     

Hydraulic lift and its influence on the water content of the rhizosphere: an example from sugar maple,Acer saccharum

Hydraulic lift, the transport of water from deep in the soil through plant root systems into the drier upper soil layers, has been demonstrated in several woody plant species. Here the volume of water involved in hydraulic lift by a mature sugar maple tree is estimated. Twenty-four intact soil cores were collected from the vicinity of a sugar maple tree at the same positions at which thermocouple psychrometers had been placed. Desorption measurements were made on the soil cores and the data were fitted to the Campbell relation for soil matric potential versus soil water content . The psychrometer data were filtered to obtain the diurnal component contributed by hydraulic lift. The diurnal component in was combined with the Campbell relation for each soil core to obtain the increase in soil water content due to hydraulic lift. The additional water contents were numerically integrated to obtain a volume of 102±54 1 of water which was hydraulically lifted each night. The volume of hydraulically lifted water (HLW) is sufficiently great that in ecosystems where hydraulic lift occurs it should be included in models for calculating the water balance. However, a previous analysis of the stable hydrogen isotope composition (D) of water in understory plants around trees conducting hydraulic lift implies a much greater volume of HLW than that calculated from the analysis performed above. To reconcile these differences, it is hypothesized that some understory plants preferentially extract HLW due to its higher matric potential and that the proportion of this water source within the xylem sap of at least some understory plants that use HLW was so great that the roots of these plants must therefore be in close proximity to the tree roots from which the HLW comes. The results of this study have implications for studies of plant competition where positive associations may exist as well as for ion uptake, nutrient cycling and the design of agroforestry systems.     

白刺沙包浅层土壤水分动态及其对不同降雨量的响应

以乌兰布和沙漠典型白刺沙包为研究对象,使用EC-5土壤水分传感器对其浅层(0—50 cm)土壤水分进行长期连续监测,分析了白刺沙包不同深度土层对不同降雨量的响应及整个生长季的土壤水分动态特征。结果表明:降雨是乌兰布和沙漠白刺沙包土壤水分的最重要补给源,降雨量大小是影响浅层土壤水分补给深度的决定因素。小于10 mm的降雨完全被表层(0—10 cm)土壤吸收,无法补给10 cm以下土壤水分;10—20 mm的降雨对土壤水分的补给深度达到20 cm;20—30 mm的降雨对土壤水分的补给深度达到40 cm,大于30 mm的降雨补给深度可达到50 cm,甚至更深土层。在研究区降雨量以小于20 mm降雨为主的情况下,20 cm以下土层土壤水分逐步恶化,久之将有利于浅根系草本植物的生长,不利于白刺的生长繁殖。因此,这种降雨格局将对浅层土壤水分及植被演替产生重要影响。     

多枝柽柳幼苗根系形态及生物量对不同灌溉处理的响应

多枝柽柳(Tamarix ramosissima)是塔里木河下游荒漠河岸林中的优势灌木, 对荒漠河岸植被群落的稳定起着重要作用。该文通过研究多枝柽柳幼苗根系形态对不同灌溉处理的响应, 分析人工水分干扰对多枝柽柳幼苗根系生长的影响。实验设计了侧渗分层和地表灌溉两种给水方式和高灌(50 L∙株 ^-1)、中灌(25 L∙株 ^-1)、低灌(12.5 L∙株 ^-1)三个给水水平, 并在整个生长季节监测每个植株的生物量及根系形态参数。结果显示: 与地表灌溉比较, 侧渗分层的灌溉方式显著提高了细根(0.5 mm < d < 2 mm)长、细根表面积和根系生物量, 并使根系生长至160 cm深度的土层, 大于地表灌溉深度(80-100 cm); 侧渗分层灌溉+高灌的组合促进根系生长的效果最显著(p < 0.05); 侧渗分层灌溉方式下总细根(d < 2 mm)的比根长随着给水量的增加显著增大, 而地表灌溉下比根长无显著变化; 侧渗分层灌溉方式下根冠比总体小于地表灌溉方式, 即侧渗分层灌溉使多枝柽柳地上部分发育较好。因此, 侧渗分层灌溉方式有显著促进多枝柽柳幼苗在生长早期快速发育的效果。     

绿洲前沿地区多枝柽柳水分关系的特征及灌溉的影响

 对塔克拉玛干沙漠南缘策勒绿洲前沿地区多枝柽柳（Tamarix ramosissima）进行了研究,生长季节和灌溉前后水分关系的特征表明：多枝柽柳在生长季节保持了较高较稳定的清晨水势,植物能够平衡白天水分损失造成的水分亏缺,水分恢复状况良好。环境气候变化对渗透势等水分参数的影响不及植物水分恢复状况对它们的影响。除了个别数据,多枝柽柳水势和渗透势的降低幅度很小,更像是生长过程的结果,植物水分胁迫的特征并不明显。对比水分生理上对干旱胁迫的适应——渗透势的迅速降低和质外体水含量的增加,膨压消失点相对含水量(RWCp)和渗透势差值(ΔΠ)等参数显示的生理特征表明,植物更加依靠稳定充足的水分供应来适应生长环境中极端的高水压差和与此相应的高大气蒸发要求。与此适应,植物和地下水发生了联系,并且一次性灌溉对植物水分状况的恢复基本没有帮助,保持地下水位稳定在根系的可吸收范围内成为保护绿洲前沿多枝柽柳长期存在的关键。多枝柽柳的水分特征属于中生植物的范畴,对极端环境的适应属于躲避的类型。     

Influence of root density on the critical soil water potential

Estimation of root water uptake in crops is important for making many other agricultural predictions. This estimation often involves two assumptions: (1) that a critical soil water potential exists which is constant for a given combination of soil and crop and which does not depend on root length density, and (2) that the local root water uptake at given soil water potential is proportional to root length density. Recent results of both mathematical modeling and computer tomography show that these assumptions may not be valid when the soil water potential is averaged over a volume of soil containing roots. We tested these assumptions for plants with distinctly different root systems. Root water uptake rates and the critical soil water potential values were determined in several adjacent soil layers for horse bean (Vicia faba) and oat (Avena sativa) grown in lysimeters, and for field-grown cotton (Gossypium L.), maize (Zea mays) and alfalfa (Medicago sativa L.) crops. Root water uptake was calculated from the water balance of each layer in lysimeters. Water uptake rate was proportional to root length density at high soil water potentials, for both horse bean and oat plants, but root water uptake did not depend on root density for horse bean at potentials lower than −25 kPa. We observed a linear dependency of a critical soil water potential on the logarithm of root length density for all plants studied. Soil texture modified the critical water potential values, but not the linearity of the relationship. B E Clothier Section editor