黑河下游荒漠河岸植物根系水力再分配及其生态水文效应
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摘要
在水分胁迫条件下,根系的“开源”和气孔的“节流”作用是植物延迟脱水的主要机理。水力再分配(HR)是一种有助于根系增加吸水以延缓脱水的可能机理,即在土壤水势梯度驱动下经由根系将土壤水分由供应充足部位转移到严重缺水根际环境的一种被动运输过程,主要包括水力提升(HL)、水力下传(HD)、侧向再分配(LR)、叶片吸收(FU)和组织脱水(TD)5种模式。目前,已在全球不同生态系统约120种植物中发现根系HR,并对其大小及生态水文效应开展了广泛而深刻的讨论。国内对根系HR的研究起步较晚,手段单一,基本停留在HL这一层面,对其生态水文效应的研究更是鲜见报道。因此,本文采用热比率法、十壤水分观测及其不同尺度的植物水分生理观测等探究了黑河下游荒漠河岸植物胡杨(Populus euphratica Oliv.)和柽柳(这里指多枝柽柳Tamarix ramosissima Ledeb.)根系HR及其生态水文效应。
(1)水力再分配过程
胡杨根系不仅具有HL,降水后也能将表面湿土层吸收的水分向下运输,即HD。同时,在河道灌水后,根系也能将河道内湿土中吸收的水分运输至距离河道较远的干土中,即LR。有趣的是,LR过程中侧根不同深度存在双向液流,且树干液流速率也明显减小,说明这一过程可能还受到树干调节,可能的机理是根木质部的径向分区(Radial sectoring),即侧根吸收的水分向上运输至茎木质部后沿着心材作螺旋运动而将一部分水分向下运输至另一侧的根中。尽管柽柳侧根液流未出现负向流动,但有两个证据表明柽柳根系HR确实存在：(i)根际区土壤含水量的昼夜波动；(ii)稳定同位素确认的水分主要来源于地下水和浅层土壤水。柽柳根际区大量不定根的存在,说明柽柳HR的路径可能不是侧根而是不定根。
(2)水力再分配的大小及影响因素
胡杨根系水力再分配的数量小于柽柳。对于胡杨,基于侧根负向液流计算的生长季根系HR大小在0.16～0.26mmd-1之间变化,平均为0.21mmd-1。对于柽柳,基于土壤体积含水量变化估算的根系HR大小在0.14～1.02mmd-1之间变化,平均为0.48mmd-1。相关分析和逐步回归分析表明：根系HR的大小除受蒸腾拉力控制外,还与气候因子和土壤水分有效性显著相关,其中：HR与水汽压差、土壤温度、土壤含水量呈显著的正相关,而与相对湿度呈显著的负相关。
(3)水力再分配的水文效应
首先,HR能够调节根际区土壤水分平衡。以柽柳为例,在干季(7月14-23日),表层80cm土壤内,每天夜间HR(5.42mm)补充了大约71.00%的白天损耗(7.59mm)。由于HR的存在,土壤储水量减小比自然条件下减少到相同水平至少要提前约4天。其次,水力再分配通过对土壤水分有效性的调节而间接地影响着植物蒸腾。生长季胡杨根系HR对蒸腾的贡献在14.60～113.04%之间,平均为38.75%；柽柳根系HR对蒸腾的贡献在10.46～53.33%之间,平均为19.44%。最后,水力再分配也影响着林分水量平衡。2012年生长季,胡杨林分HR(123.00mm)对蒸散发(313.00mm)的贡献约为37.00%,柽柳林地HR(117.20mm)对蒸散发(607.27mm)的贡献约为19.00%。另外,林分蒸散发差异也影响到局地小气候。典型日(7月12-21日),日间胡杨林地温度比柽柳林地最大高约4℃,相对湿度低约10.00%,水汽压差高约1.60kPa。
(4)水力再分配的生态效应
根系水力再分配最直接的表现是维持浅层土壤中细根有效性,进而能够对干旱区降水脉冲作出快速响应。换言之,HD可以通过深层土壤水补给影响整个生态系统水分平衡。一次降水事件后,水分通过HR向深层土壤的运移要比入渗或优势流更快。如果不存在HR,深层土壤的水分补给将更低。根系水力再分配通过增加植物水分吸收而影响着植物叶片的气体交换参数。另外,根系水力再分配也影响植物群落的组成与格局,这有待进一步证实。
与传统观点对于水分利用策略的认识不同,本文认为：在地下水埋深较浅时,柽柳根系能够通过HR吸收深层丰富的水分和浅层富集的养分,以保持气孔开放维持高的蒸腾速率,这使得它们每天的光合产物积累和生长速率大于胡杨,即在水分供应充足条件下柽柳为耗水植物。在水分胁迫条件下(干季),叶水势下降,气孔关闭并在厚角质层和浓密的表皮保护下蒸腾耗水降到最低,即由耗水植物变为节水植物。但是,由于干旱时叶子(同化枝)生长减缓,过量的光合产物被运到根部,导致根的更快生长,再向湿润土壤深入吸收更多的水分。如此往复,使柽柳根与地上部比例进一步增大。因此,像柽柳这样的耗水性旱生植物要比胡杨这样的节水性旱生植物更加耐旱,这可能就是柽柳在中亚广泛分布,而在美国西南部河流沿岸疯狂生长的原因。
In desert ecosystems, the mainly mechanisms of plants delay dehydration to resist droughts is to increase water income via roots and reduce it expenditure via stoma of leaf. To tolerate drought during dry season, desert also riparian plants undergo hydraulic redistribution (HR), the passive movement of water between different soil parts via plant root systems, driven by water potential gradients in the soil-plant interface. Patterns of HR include hydraulic lift (HL), hydraulic descent (HD), lateral redistribution (LR), foliar uptake (FU) and tissue dehydration (TD). Now, HR has been described in approximately120species that involves a wide variety of ecosystems and a wide range of life forms. Apart from its magnitude, the potential ecohydrologic consequences of HR have attracted recent attention. However, the study status in China is started late, single means and the content focus on HL and the rarely reported refer to ecohydrologic consequences of HR. The objectives of this study were (1) to search for the evidence, quantify the magitude and discuss the ecohydrologic consequences of that the roots of two desert riparian plants, Populus euphratica Oliv. and Tamarix ramosissima Ledeb., carry out HR. To demonstrate HR, we present data on patterns of sap flow in the stems or branches and lateral roots of those two desert riparian species and soil volumetric moisture content where these species grow, and the datas of plant water physiology with different scales also been provided.
(1) Patterns of hydraulic redistribution
We not only confirm the previous knowledge on HL of P. euphratica, but also the water can transport from moist topsoil to dry subsoil after rain, i.e. HD. In addition, water also moved from lateral moist soil layer to opposite dry soil layer, i.e. LR. Interestingly, we observed bidirectional flow in the lateral roots of P. euphratica during the time of lateral redistribution, and it may be mediated by stem tissues and, by inference, the radial sectoring in the xylem that the stem base seems in connection with the root xylem. The water absorbing from the wet side adjacent to river was transported to the stem; the circumferential movement of sap flow around the heartwood becomes available to the downstream flow. Although no direct evidence indicated reverse sap flow of lateral roots and associated HR in T. ramosissima, several factors indicate that HR is occurring:(1) diel fluctuations of volumetric moisture content in the upper soil layer and (ii) the identification of primary water sources as groundwater and vadose zone water through stable isotope studies. The reason of HR not occurred in lateral roots of T. ramosissima was potential nocturnal transpiration suppression. Thus, we inferred that HR occurs in T. ramosissima via adventitious roots rather than via lateral roots.
(2) Magitudes and controlling factors of hydraulic redistribution
For P. euphratica. the magnitude of HR based on nagitive sap flow in lateral roots was ranged from0.16mm d-1to0.26mm d-1with an average of0.21mm d-1. For T. ramosissima, the magnitude of HR based on diurnal fluctuations of soil volumetric moisture content at20cm to60cm depths was ranged ranged from0.14mm d-1to1.02mm d-1with an average of0.48mm d-1during the growing season, which was far less than that in2011. The correlation and stepwise regression analysis demonstrated that apart from the transpiration tension, the climate and soil moisture availability can accounted for the variation of HR. Specifically, HR was significantly positive correlation with vapor pressure deficit, soil temperature and soil moisture content, instead negative correlation with relative humidity.
(3) Hydrological effects of hydraulic redistribution
The effect of HR on soil moisture balance is positive. For example, in T. ramosissima stand, approximately71.00%of the water removed daily (7.59mm) from the upper80cm of soil was replaced by nocturnal hydraulic redistribution (5.42mm) during dry season (14to23July). If hydraulic redistribution had not occurred, the soil water content observed at the end of the10-day monitoring period would have been attained about4days sooner. Because of the increase of soil moisture, the transpiration was also increased that indirectly induced by HR. For P. euphratica. hydraulic redistribution increased transpiration ranged from14.60%to113.04%with an average of38.75%during the growing season. For T. ramosissima, hydraulic redistribution increased transpiration ranged from10.46%to53.33%with an average of19.44%. In the same way, the effect of HR on community water balance is positive. In2012, approximately37.00%of evapotranspiration (313.00mm) was replaced by nocturnal hydraulic redistribution (123.00mm) during dry season for P. euphratica stand. Accordingly, approximately19%of evapotranspiration (607.27mm) was replaced by nocturnal hydraulic redistribution (117.20mm) during dry season for T. ramosissima stand. In addition, the difference of evapotranspiration also affects the microclimate among stands, that the temperature and vapor pressure deficit for P. euphratica stand was maximal higher4℃and1.60kPa than T. ramosissima stand, respectively, instead relative humidity for the former maximal lower10%than the latter during the typical days (12to21July).
(4) Ecological effect of hydraulic redistribution
The directly ecological effect of hydraulic redistribution on plants were to maintain the effectiveness of fine roots in the shallow soil, that made it can quickly reaponse to precipitation pulse in arid area. HD, on the other hand, can affect overall ecosystem water budgets by increasing deep soil water recharge. Water movement to deep soil layers after a rain event is faster through HR than via infiltration or preferential flow and, in the absence of HR. water recharge of deep soil layers would be very low. And then gas exchange parameters of leaf were affected by increased water absorbing via fine roots in the shallow soil. Finally, HR may also have effects on plant community composition and structure, that further investigation is needed to substantiate this.
In contrast to the conventional view on water use strategies of plants, we thought the roots of T. ramosissima can simultaneously absorb plentiful water in deep soil layer and nutrient in shallow soil layer throught HR, which made the leaves of T. ramosissima can keep stoma open and maintain higher transpiration rate and photosynthetic product accumulation than P. euphratica. When soil water is limited, transpiration reduced rapidly to a minimum under the protection of thickly cutin layer and skin, along with descend of the leaf water potential and closure of stoma, which suggest the transform of T. ramosissima from water spender to saver. However, because the growth decreases of leaf or assimilating shoots, the excessive amounts of photosynthetic products was transport to the roots that lead to the faster growth and then more water absorbing in deep soil layer. So given enough time, the proportion of roots and overground part will increase further. Therefore, xerophytes of water-consuming as T. ramosissima is more drought tolerant than water-saving like P. euphratica. it could be the reasons that widely distributed in central Asia and crazy grow along rivers in southwestern United States.
(1)水力再分配过程
胡杨根系不仅具有HL,降水后也能将表面湿土层吸收的水分向下运输,即HD。同时,在河道灌水后,根系也能将河道内湿土中吸收的水分运输至距离河道较远的干土中,即LR。有趣的是,LR过程中侧根不同深度存在双向液流,且树干液流速率也明显减小,说明这一过程可能还受到树干调节,可能的机理是根木质部的径向分区(Radial sectoring),即侧根吸收的水分向上运输至茎木质部后沿着心材作螺旋运动而将一部分水分向下运输至另一侧的根中。尽管柽柳侧根液流未出现负向流动,但有两个证据表明柽柳根系HR确实存在：(i)根际区土壤含水量的昼夜波动；(ii)稳定同位素确认的水分主要来源于地下水和浅层土壤水。柽柳根际区大量不定根的存在,说明柽柳HR的路径可能不是侧根而是不定根。
(2)水力再分配的大小及影响因素
胡杨根系水力再分配的数量小于柽柳。对于胡杨,基于侧根负向液流计算的生长季根系HR大小在0.16～0.26mmd-1之间变化,平均为0.21mmd-1。对于柽柳,基于土壤体积含水量变化估算的根系HR大小在0.14～1.02mmd-1之间变化,平均为0.48mmd-1。相关分析和逐步回归分析表明：根系HR的大小除受蒸腾拉力控制外,还与气候因子和土壤水分有效性显著相关,其中：HR与水汽压差、土壤温度、土壤含水量呈显著的正相关,而与相对湿度呈显著的负相关。
(3)水力再分配的水文效应
首先,HR能够调节根际区土壤水分平衡。以柽柳为例,在干季(7月14-23日),表层80cm土壤内,每天夜间HR(5.42mm)补充了大约71.00%的白天损耗(7.59mm)。由于HR的存在,土壤储水量减小比自然条件下减少到相同水平至少要提前约4天。其次,水力再分配通过对土壤水分有效性的调节而间接地影响着植物蒸腾。生长季胡杨根系HR对蒸腾的贡献在14.60～113.04%之间,平均为38.75%；柽柳根系HR对蒸腾的贡献在10.46～53.33%之间,平均为19.44%。最后,水力再分配也影响着林分水量平衡。2012年生长季,胡杨林分HR(123.00mm)对蒸散发(313.00mm)的贡献约为37.00%,柽柳林地HR(117.20mm)对蒸散发(607.27mm)的贡献约为19.00%。另外,林分蒸散发差异也影响到局地小气候。典型日(7月12-21日),日间胡杨林地温度比柽柳林地最大高约4℃,相对湿度低约10.00%,水汽压差高约1.60kPa。
(4)水力再分配的生态效应
根系水力再分配最直接的表现是维持浅层土壤中细根有效性,进而能够对干旱区降水脉冲作出快速响应。换言之,HD可以通过深层土壤水补给影响整个生态系统水分平衡。一次降水事件后,水分通过HR向深层土壤的运移要比入渗或优势流更快。如果不存在HR,深层土壤的水分补给将更低。根系水力再分配通过增加植物水分吸收而影响着植物叶片的气体交换参数。另外,根系水力再分配也影响植物群落的组成与格局,这有待进一步证实。
与传统观点对于水分利用策略的认识不同,本文认为：在地下水埋深较浅时,柽柳根系能够通过HR吸收深层丰富的水分和浅层富集的养分,以保持气孔开放维持高的蒸腾速率,这使得它们每天的光合产物积累和生长速率大于胡杨,即在水分供应充足条件下柽柳为耗水植物。在水分胁迫条件下(干季),叶水势下降,气孔关闭并在厚角质层和浓密的表皮保护下蒸腾耗水降到最低,即由耗水植物变为节水植物。但是,由于干旱时叶子(同化枝)生长减缓,过量的光合产物被运到根部,导致根的更快生长,再向湿润土壤深入吸收更多的水分。如此往复,使柽柳根与地上部比例进一步增大。因此,像柽柳这样的耗水性旱生植物要比胡杨这样的节水性旱生植物更加耐旱,这可能就是柽柳在中亚广泛分布,而在美国西南部河流沿岸疯狂生长的原因。
In desert ecosystems, the mainly mechanisms of plants delay dehydration to resist droughts is to increase water income via roots and reduce it expenditure via stoma of leaf. To tolerate drought during dry season, desert also riparian plants undergo hydraulic redistribution (HR), the passive movement of water between different soil parts via plant root systems, driven by water potential gradients in the soil-plant interface. Patterns of HR include hydraulic lift (HL), hydraulic descent (HD), lateral redistribution (LR), foliar uptake (FU) and tissue dehydration (TD). Now, HR has been described in approximately120species that involves a wide variety of ecosystems and a wide range of life forms. Apart from its magnitude, the potential ecohydrologic consequences of HR have attracted recent attention. However, the study status in China is started late, single means and the content focus on HL and the rarely reported refer to ecohydrologic consequences of HR. The objectives of this study were (1) to search for the evidence, quantify the magitude and discuss the ecohydrologic consequences of that the roots of two desert riparian plants, Populus euphratica Oliv. and Tamarix ramosissima Ledeb., carry out HR. To demonstrate HR, we present data on patterns of sap flow in the stems or branches and lateral roots of those two desert riparian species and soil volumetric moisture content where these species grow, and the datas of plant water physiology with different scales also been provided.
(1) Patterns of hydraulic redistribution
We not only confirm the previous knowledge on HL of P. euphratica, but also the water can transport from moist topsoil to dry subsoil after rain, i.e. HD. In addition, water also moved from lateral moist soil layer to opposite dry soil layer, i.e. LR. Interestingly, we observed bidirectional flow in the lateral roots of P. euphratica during the time of lateral redistribution, and it may be mediated by stem tissues and, by inference, the radial sectoring in the xylem that the stem base seems in connection with the root xylem. The water absorbing from the wet side adjacent to river was transported to the stem; the circumferential movement of sap flow around the heartwood becomes available to the downstream flow. Although no direct evidence indicated reverse sap flow of lateral roots and associated HR in T. ramosissima, several factors indicate that HR is occurring:(1) diel fluctuations of volumetric moisture content in the upper soil layer and (ii) the identification of primary water sources as groundwater and vadose zone water through stable isotope studies. The reason of HR not occurred in lateral roots of T. ramosissima was potential nocturnal transpiration suppression. Thus, we inferred that HR occurs in T. ramosissima via adventitious roots rather than via lateral roots.
(2) Magitudes and controlling factors of hydraulic redistribution
For P. euphratica. the magnitude of HR based on nagitive sap flow in lateral roots was ranged from0.16mm d-1to0.26mm d-1with an average of0.21mm d-1. For T. ramosissima, the magnitude of HR based on diurnal fluctuations of soil volumetric moisture content at20cm to60cm depths was ranged ranged from0.14mm d-1to1.02mm d-1with an average of0.48mm d-1during the growing season, which was far less than that in2011. The correlation and stepwise regression analysis demonstrated that apart from the transpiration tension, the climate and soil moisture availability can accounted for the variation of HR. Specifically, HR was significantly positive correlation with vapor pressure deficit, soil temperature and soil moisture content, instead negative correlation with relative humidity.
(3) Hydrological effects of hydraulic redistribution
The effect of HR on soil moisture balance is positive. For example, in T. ramosissima stand, approximately71.00%of the water removed daily (7.59mm) from the upper80cm of soil was replaced by nocturnal hydraulic redistribution (5.42mm) during dry season (14to23July). If hydraulic redistribution had not occurred, the soil water content observed at the end of the10-day monitoring period would have been attained about4days sooner. Because of the increase of soil moisture, the transpiration was also increased that indirectly induced by HR. For P. euphratica. hydraulic redistribution increased transpiration ranged from14.60%to113.04%with an average of38.75%during the growing season. For T. ramosissima, hydraulic redistribution increased transpiration ranged from10.46%to53.33%with an average of19.44%. In the same way, the effect of HR on community water balance is positive. In2012, approximately37.00%of evapotranspiration (313.00mm) was replaced by nocturnal hydraulic redistribution (123.00mm) during dry season for P. euphratica stand. Accordingly, approximately19%of evapotranspiration (607.27mm) was replaced by nocturnal hydraulic redistribution (117.20mm) during dry season for T. ramosissima stand. In addition, the difference of evapotranspiration also affects the microclimate among stands, that the temperature and vapor pressure deficit for P. euphratica stand was maximal higher4℃and1.60kPa than T. ramosissima stand, respectively, instead relative humidity for the former maximal lower10%than the latter during the typical days (12to21July).
(4) Ecological effect of hydraulic redistribution
The directly ecological effect of hydraulic redistribution on plants were to maintain the effectiveness of fine roots in the shallow soil, that made it can quickly reaponse to precipitation pulse in arid area. HD, on the other hand, can affect overall ecosystem water budgets by increasing deep soil water recharge. Water movement to deep soil layers after a rain event is faster through HR than via infiltration or preferential flow and, in the absence of HR. water recharge of deep soil layers would be very low. And then gas exchange parameters of leaf were affected by increased water absorbing via fine roots in the shallow soil. Finally, HR may also have effects on plant community composition and structure, that further investigation is needed to substantiate this.
In contrast to the conventional view on water use strategies of plants, we thought the roots of T. ramosissima can simultaneously absorb plentiful water in deep soil layer and nutrient in shallow soil layer throught HR, which made the leaves of T. ramosissima can keep stoma open and maintain higher transpiration rate and photosynthetic product accumulation than P. euphratica. When soil water is limited, transpiration reduced rapidly to a minimum under the protection of thickly cutin layer and skin, along with descend of the leaf water potential and closure of stoma, which suggest the transform of T. ramosissima from water spender to saver. However, because the growth decreases of leaf or assimilating shoots, the excessive amounts of photosynthetic products was transport to the roots that lead to the faster growth and then more water absorbing in deep soil layer. So given enough time, the proportion of roots and overground part will increase further. Therefore, xerophytes of water-consuming as T. ramosissima is more drought tolerant than water-saving like P. euphratica. it could be the reasons that widely distributed in central Asia and crazy grow along rivers in southwestern United States.
引文
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