Hydraulic characteristics and embolism repair of Populus alba × P. glandulosa after drought stress and rehydration
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摘要:目的 研究植物遭受不同程度干旱胁迫时，其水力学特性的变化及响应，以及复水后植物栓塞修复能力，为植物应对干旱环境的能力提供水力学依据。方法 以84K杨扦插苗为研究对象，进行渐进的控水处理，根据植株形态特征的变化，分别控水至植株叶面积停止生长、整株萎蔫、50%的叶片死亡及全部叶片死亡4个阶段，而后各阶段植株均进行复水至新生叶片长出。分别在控水和复水处理完成后，测定各阶段植株的木质部水势、叶水势、栓塞脆弱性、枝条导水率及栓塞程度(PLC)等水力学特征，同时测定导管直径、导管连接度及导管抗垮塌能力等木质部解剖结构特征。结果 随着干旱胁迫程度加剧，84K杨叶水势及木质部水势均降低，栓塞加剧，栓塞脆弱性减小，木质部导管直径较对照组显著减小，导管连接度及导管抗垮塌能力显著增大。当植株有50%的叶片死亡时，茎的PLC为44%，当叶片全部死亡时，茎的PLC达65%。复水10 ~ 24 d后，各干旱阶段植株木质部水势及叶水势均恢复至对照组水平，茎的导水率均有所增加，栓塞程度均降低，当茎的PLC恢复至28% ~ 37%时，植株顶端重新展开了3片新叶。叶片全部死亡的植株复水至长出新叶时，茎的PLC显著减少，但植株直径并未增大，即复水阶段没有新生导管参与导水过程，表明茎导水率的恢复是由于发生了栓塞修复。结论 干旱胁迫会对植物水力特性造成不利影响，但植物也会通过改变木质部结构特征来适应环境。植物在维持叶片存活与茎导水能力之间存在一定的权衡，干旱胁迫下会牺牲叶片来维持茎的导水率。但当干旱胁迫解除后，植物的水力学特性也能得到恢复，即使整株叶片死亡的植物，复水后仍能恢复生长，叶片死亡并不能作为判断植物死亡的指标。植物恢复生长与茎导水率恢复之间存在很强的相关性，栓塞能否修复是植物经历干旱后能否存活的主要因素。Abstract:Objective The aim of this study was to explore the changes and responses of hydraulic characteristics of plants under different levels of drought stress, and their ability to repair embolism after rehydration, thereby providing theoretical hydraulic evidences for plants to adapt to drought conditions and their ability to recover after drought.Method 84K poplar (Populus alba × P. glandulosa) cuttings were potted and subjected to a progressive drought. According to the changes of plant morphological characteristics, water was controlled to four stages, i.e. the cessation of leaf expansion, whole plant wilting, 50% leaf mortality and 100% leaf mortality. Then, all plants of each stage were rewatered until new leaves appeared. After water control and rehydration treatment, hydraulic characteristics, such as xylem water potential, leaf water potential, vulnerability to xylem cavitation, stem hydraulic conductivity, the percentage loss of xylem conductance, and xylem anatomy such as the vessel diameter, contact fraction and vessel implosion resistance were measured at each stage.Result With the intensification of drought stress, leaf water potential and xylem water potential decreased, and the embolism increased, but xylem vulnerability to cavitation increased. Also, compared with the control group, the vessel diameter decreased significantly, and the contact fraction and vessel implosion resistance increased significantly. At the stage of 50% leaf mortality, the percentage loss of hydraulic conductivity (PLC) of stem was 44%, and reached 65% at the stage of 100% leaf mortality. After rehydration for 10−24 days, the xylem water potential and leaf water potential in each stage returned to the level of control group, and the stem hydraulic conductivity increased with the PLC decreased. When the PLC of stem restored to 28%−37%, three new leaves spread out. After the plants with 100% leaf mortality were rehydrated until the emergence of new leaves, the PLC of stem was significantly reduced, but the plant diameter did not increase, which showed no new conduits participated in the water transportation during the rehydration, indicating that the restoration of stem hydraulic conductivity may result from embolism repair.Conclusion Drought stress can adversely affect the hydraulic characteristics of plants, but plants can also adapt to the environment by changing the structure of their xylem. There is a certain trade-off between the plant’s ability to maintain leaf survival and transport water, and the leaves may be sacrificed to maintain the water conductivity of stem under drought stress. However, when drought stress is released, the hydraulic characteristics of the plants can also be restored. Even if plants without survival leaves can resume growth after rehydration, leaf death cannot be used as an indicator of plant death. There is a strong correlation between the recovery of plant growth and the restoration of stem hydraulic conductivity. Embolism repair may be the main reason for the restoration of stem hydraulic conductivity.
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图 2 控水及复水处理下植株直径增长
箭头表示复水时间。误差线代表 ± 标准误。Arrow represents rehydration time point, error bars represent ± SE.
Figure 2. Diameter growth of plants under drought stressand rehydration treatment
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图 3 84K杨不同控水阶段及复水后木质部水势和叶水势
不同小写字母表示控水或复水处理下不同阶段间差异性显著，不同大写字母表示同一阶段复水前后的差异性显著(P < 0.05)。下同。Different lowercase letters indicate significant differences in varied stages under drought stress or rehydration, and different capital letters indicate significant difference before and after rehydration at the same stage (P < 0.05). The same below.
Figure 3. Xylem water potential and leaf water potential of 84K poplar at different stages under drought stress and rehydration
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图 4 84K杨不同控水阶段及复水后茎的PLC
Figure 4. PLC of 84K poplar stem at different stages under drought stress and rehydration
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图 5 84K杨不同控水阶段的枝条栓塞脆弱性曲线
Figure 5. Vulnerability curves of 84K poplar at different drought stages
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图 6 84K杨不同控水阶段及复水后的栓塞脆弱性曲线
Figure 6. Vulnerability curves of 84K poplar under drought stress and rehydration
表 1 84K杨不同控水阶段及复水后栓塞脆弱性(P50)及最大导水率
Table 1 P50 and maximum hydraulic conductivity of 84K poplar under drought stress and rehydration
阶段 Stage 栓塞脆弱性 Embolism vulnerability (P50)/MPa 最大导水率 Maximum hydraulic conductivity (Kmax)/(kg·m·MPa−1·s−1) 控水 Drought stress 复水 Rehydration 控水 Drought stress 复水 Rehydration 0 −1.65 ± 0.048c 9.63 × 10−5 ± 6.60 × 10−6a 1 −1.83 ± 0.022Ab −1.84 ± 0.040Ac 7.70 × 10−5 ± 1.89 × 10−6Aab 5.88 × 10−5 ± 8.02 × 10−6Ab 2 −2.21 ± 0.029Aa −2.09 ± 0.026Ab 4.34 × 10−5 ± 1.66 × 10−6Abc 4.65 × 10−5 ± 8.04 × 10−6Abc 3 −2.23 ± 0.034Aa −2.08 ± 0.087Ab 4.14 × 10−5 ± 9.82 × 10−6Abc 3.60 × 10−5 ± 5.58 × 10−6Ac 4 −2.32 ± 0.034Aa −2.24 ± 0.017Aa 2.76 × 10−5 ± 5.32 × 10−6Ac 3.02 × 10−5 ± 5.28 × 10−6Ac 注：表中数据为平均值 ± 标准误。不同小写字母表示控水或复水下不同阶段间差异性显著，不同大写字母表示同一阶段复水前后的差异性显著(P < 0.05)。下同。Notes: data are mean ± SE. Different lowercase letters indicate significant differences in baried stages under drought stress or rehydration，and different capital letters indicate significant differences before and after rehydration at the same stage (P < 0.05). The same below. 
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表 2 84K杨复水前后木质部结构特征
Table 2 Xylem structural characteristics of 84K poplar under drought stress and rehydration
阶段 Stage 导管直径 Vessel diameter/µm 导管连接度 Vessel contact fraction 导管抗垮塌能力 Vessel implosion resistance 控水 Drought stress 复水 Rehydration 控水 Drought stress 复水 Rehydration 控水 Drought stress 复水 Rehydration 0 37.34 ± 0.15a 0.03 ± 0.001a 0.04 ± 0.003c 1 35.49 ± 0.06Ab 35.63 ± 0.18Ab 0.05 ± 0.003Ab 0.05 ± 0.004Ab 0.06 ± 0.005Aa 0.06 ± 0.003Aa 2 34.97 ± 0.07Ac 34.47 ± 0.16Ac 0.05 ± 0.004Ab 0.05 ± 0.002Ab 0.05 ± 0.001Ab 0.05 ± 0.002Ab 3 34.77 ± 0.20Ac 34.51 ± 0.03Ac 0.05 ± 0.003Ab 0.04 ± 0.005Ab 0.05 ± 0.001Ab 0.05 ± 0.002Ab 4 34.80 ± 0.31Ac 34.85 ± 0.30Ac 0.04 ± 0.005Ab 0.05 ± 0.002Ab 0.05 ± 0.001Ab 0.05 ± 0.003Ab
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