黄土丘陵沟壑区优势植物对不同侵蚀环境的适应研究
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摘要
黄土高原严重的土壤侵蚀，造成了该区梁峁起伏、地形破碎、立地环境复杂多变，使得水、热等生态因子的空间分布存在异质性，形成了多样的侵蚀环境。植物随着生长环境的改变，在群落组成、生殖策略、形态结构、生理特性上不断的调整来适应相应的环境。本文通过陕北黄土丘陵沟壑区3个植被带（森林带、森林草原带、草原带）坡沟系统中5种不同侵蚀环境下（阳沟坡、阳峁坡、峁顶、阴峁坡、阴沟坡）植物群落组成的调查，分析群落中各物种的生态位宽度和重要值，据此筛选出不同侵蚀环境下的优势物种。研究了九种优势植物（猪毛蒿、铁杆蒿、茭蒿、白羊草、长芒草、芦苇、达乌里胡枝子、杠柳、狼牙刺）在不同侵蚀环境下的繁殖、形态、解剖和生理响应特征，并从中选取了4种枝系构型植物（聚丛型铁杆蒿、疏丛型茭蒿、簇丛型白羊草、主杆型狼牙刺），研究了其对坡面微环境的影响。主要研究结果如下：
（1）由森林带向草原带过渡，植物群落结构变的简单，物种多样性减小，但物种个数反而有增加趋势；中生植物、高位芽植物减少，旱生植物、隐芽植物增加。坡沟系统不同侵蚀环境下物种均匀度差异不显著，但阴坡群落物种多样性和丰富度较阳坡提高；阳沟坡旱生植物和隐芽植物比例大；而峁顶一年生植物比例增大。从阳沟坡至阴沟坡随着侵蚀程度的减小，物种数量和密度增加，但各物种生态位宽度和重要值减小。旱生和旱中生植物在黄土丘陵沟壑区有着广泛的分布。
（2）同种植物单个花序（花）重、单个种子重量表现出一定的遗传稳定性，在坡沟系统不同侵蚀环境下差异不明显。阳沟坡和阴沟坡植物有性生殖投入低，但该侵蚀环境下隐芽植物比例大，使得其上物种主要以芽库完成地上植被更新。峁顶和阳峁坡植物有性生殖投入量大，产生大量的种子有利于植物种群的扩大。相对其他供试物种，达乌里胡枝子有性繁殖分配比例大，而两种灌木狼牙刺与杠柳则相反。在芽库季节动态上，植物返青期较大的永久性芽库存量保证了植物迅速更新，开花结实期大量季节性芽库确保了植物光合产物积累。
（3）同种植物叶片形态解剖结构在3个植被带相同坡沟侵蚀环境下变化差异性不显著，但坡沟系统不同的侵蚀环境下植物叶片形态和解剖结构变化差异明显。随着侵蚀程度的增加，叶片叶片厚度、干物质量积累、角质层厚度、栅栏组织厚度、木质部韧皮部面积比增加，叶面积、表皮细胞大小、海绵组织厚、细胞间隙减小。输送水分的木质部与植物适应环境能力关系密切。不同植物适应黄土高原半干旱的气候也有不同的策略：猪毛蒿叶具有发达贮水组织，铁杆蒿全栅型叶片，茭蒿和狼牙刺具有双栅型叶片和大的维管束鞘细胞，达乌里胡枝子叶具有发达的粘液细胞，白羊草和芦苇叶上表皮具有发达的泡状细胞，长芒草叶上表皮下多层厚壁组织，杠柳厚的角质层。综合隶属函数分析表明，叶片保护组织发达的长芒草从叶片形态与解剖角度分析对环境适应能力最强。
（4）三个植被带相同坡沟侵蚀环境下植物叶片生理特性变化未达到显著性，但坡沟系统不同侵蚀环境下植物叶片生理特性差异明显。从阴沟坡至阳沟坡随着侵蚀程度增加，细胞受伤害和叶绿素受破坏程度增加，但植物通过叶片保水力、渗透调节物质、抗逆酶活性、非酶抗氧化物质的加强来应对环境胁迫，同时减小叶片相对含水量、自由束缚水比值，以牺牲植物生长速率的代价来应对胁迫环境。所测生理指标中，与植物环境适应相关性最大的是抗坏血酸氧化酶。隶属函数综合分析表明两种灌木杠柳和狼牙刺从生理角度分析对环境适应性较强。
（5）不同构型植物拦截坡面径流沉积物和保护植冠下土体而形成不同形态的土堆。随着坡度的增加，土堆由对称堆型圆帽结构转变为不对称的台阶型结构，主杆型狼牙刺转折点在15-25°的陡坡，聚丛型铁杆蒿和簇丛型白羊草在26-35°的极陡坡，疏丛型茭蒿在≥36°的险坡。坡度和植丛顺坡基部直径影响着土堆的高度；而植丛垂直坡面的基部直径、植丛分枝数量及整个植丛的冠幅影响着土堆的面积。随着坡度的增加土堆高度总体上呈增加趋势而土堆面积趋势相反。植物拦截形成的土堆成为坡面的“营养岛”，相对于植被间表现出良好的土壤理化特性及较高的物种多样性。茭蒿以其发达且扩张的枝系、狼牙刺以其较大的植冠在形成土堆和对坡面微环境的改善方面能力较强。
（6）在繁殖策略方面，供试物种猪毛蒿、铁杆蒿、茭蒿、白羊草、杠柳趋向于r-生殖策略，长芒草、达乌里胡枝子、狼牙刺更趋向于K-生殖策略。依据植物适应环境时的形态解剖与生理特点，供试植物可分为Ⅰ：御逆型，猪毛蒿、芦苇、长芒草；Ⅱ：胁变修复型，茭蒿、白羊草、达乌里胡枝子；Ⅲ：御胁变型，铁杆蒿、狼牙刺、杠柳。综合植物繁殖、形态、解剖和生理特征，供试物种综合隶属函数值与物种重要值呈正相关关系，供试植物的侵蚀环境适应能力表现为：两种灌木狼牙刺和杠柳较强，在黄土丘陵沟壑区不同侵蚀环境有着广泛分布的铁杆蒿、达乌里胡枝子、长芒草次之，而对侵蚀环境具有选择性的白羊草、茭蒿、芦苇和猪毛蒿相对较小。
The serious soil erosion on the Loess Plateau cause the deep cut gullies, fragmentation oflandform and form a complicated landform, thus result in spatial heterogeneity of water andheat. Then, there are series of adjustments occur on plant community structure, reproductivestrategies, leaf morphological and physiological characteristics to adapt different soilenvironments. In this study, the community structures in five soil erosion environments onslope-gully system (sunny gully slope, sunny hilly slope, hilltop, shady hilly slope, and shadygully slope) on three vegetation zones (forest zone, forest-steppezone, and steppe zone) on thehilly-gully region of the Loess Plateau were examined. The species niche breadth andimportant value in community were analysed to identify dominant species at different soilerosion environments. The reprodution, morphology, anatomy, and physiology of seclectednine dominant species (Artemisia scoparia, Artemisia gmelinii, Artemisia giraldii,Bothriochloa ischcemum, Stipa bungeana, Phragmites australis, Lespedeza davurica,Periploca sepium and Sophora davidii) were studied. And, four typical shoot architecturespecies (A. gmelinii, with dense shoot architecture; A. giraldii, with expanding shootarchitecture; B. ischaemum, with a tussock-formingshoot architecture; and S. viciifolia, with amain-stem shoot architecture) was selected to analysis the shape of phytogenic mounds andits influence of mirco-environmnent on slope. The main results are as follows:
(1) Fom forest to steppe zone, plant community structures become simple, speciesdiversity decreased, and the number of species showed a slight increased with mesophytesand phanerophytes decreased, while xerophytes and cryptophytes increased. At different soilerosion environments on slope-gully system, the variation of species evenness was notsignificant. On shady slope, the species diversity and richness were relative larger than sunnyslope. The xerophyte and cryptophytes increased at sunny gully slope. Hilltop had a higher therophyte proportion than the other erosion environments. From sunny to shady gully slope,the species niche and important value in community decreased with the decreasing of erosiondegree. Xerophyte and xero-mesophytes have a wide range of distribution on the hilly-gullyregion of the Loess Plateau.
(2) For same species, single dry weight of inflorescence (flower) and seed mass showedcertain genetic stability, and no significant difference existed in different soil erosionenvironments on slope-gully system. At the sunny gully slope and shady gully slope, plantreproductive investment was low, but the high proportion of cryptophytes on theseenvironments made vegetation regeneration relying mainly on bud bank. The reproductiveinvestment of species on hilltop and sunny hilly slope were larger than the other three erosionenvironments which result in a large number of seeds and thus was beneficial to the expansionof seed dispersal plants. L. davurica had a larger reproductive allocation, while two shrubs (P.sepium and S. davidii) were relatively less. In vegetation turning-green season, the largerperennial bud bank ensure a rapid vegetation renewal. The larger seasonal bud bank onfruit-setting season ensure the accumulation of photosynthate.
(3) The leaf morphological and anatomical characteristics at same slope-gully erosionenvironment on the three vegetation zones had a slight changes, but these characteristicsshowed significant changes at different erosion environments on slope-gully system. With theincreasing of soil erosion degree, leaf thickness, dry matter content, cuticle thickness, palisadetissue thickness, xylem area/phloem area increased, while leaf area, epidermal cell size,spongy tissue thickness, intercellular space decreased. In those leaf morphological andanatomical indices, xylem area, which mediate plant water transportation, had the closestrelationship with plant environmental adaptability. Different plants have its special structureto adapt the semi-arid climate on the Loess Plateau: water storage parenchyma of A. scoparia,total-palisade tissue of A. gmelinii, bi-palisade tissue and big bundle sheath cells of A. giraldiiand S. davidii, mucilage cell of L. daurica; special bulliform cells of B. ischcemum and P.australis; multilayer sclerenchyma of S. bungeana; thick cuticle of P. sepium. From leafmorphological and anatomical characteristics, comprehensive analysis of the subordinatefunction showed that S. bungeana, which has a well developed protective tissues, had thestrongest environmental adaptability.
(4) Similar to plant morphological characteristics, plant physiological indices at sameslope-gully erosion environment on the three vegetation zones showed a slight changes, butthose characteristics presented significant changes at different erosion environments on slope-gully system. From sunny to shady gully slope，with the increasing of soil erosiondegree, the degree of cell damage and amount of chlorophyll destruction increased. But plantresistance to environmental stress by increasing leaf water-holding ability, osmotic adjustmentsubstances, stress-resistance enzymes, non-enzymatic antioxidants, meanwhile by decreasingleaf relative water content and free water/bound water. This processes are at the expense ofplant growth rate to deal with the environmental stress. Among the21measured physiologicalindices, ascorbic acid oxidase (APX) had the closest relationship with plant environmentaladaptability. From plant physiological indices, comprehensive analysis of the subordinatefunction showed that two testing shrubs, S. davidii and P. sepium, had a strong environmentaladaptability.
(5) Different shoot architecture species have different ability of preventing soil erosionand retaining sediment, thus the shape of phytogenic mound changes with slope gradientincreasing. Mound shapes changed from symmetrical mound-type structures intoasymmetrical terrace-type structures as slope gradient increased, but the change at0-15°forS. viciifolia, at26-35°for A. gmelinii and B. ischaemum, at>35°for A. giraldii. Among theplant shoot architecture parameters, plant basal diameter along the slope was significantlycorrelated with mound height, while the plant basal diameter perpendicular to the slope andtotal cross-sectional area of the stem at the plant base were associated with mound area.Mound height increased with slope for all species, while mound area showed oppositetendency. The phytogenic mounds can act as ‘resource islands’, thus improved speciesrichness and stress resistance of plant community on slope. A. giraldii with expanding shootarchitecture and S. viciifolia with its large canopy can create relatively large mounds.
(6) In reproductive strategy, K-reproductive strategyare adopted by A. scoparia, A.gmelinii, A.giraldii, B. ischcemum and P. sepium; and r-reproductive strategy by S. bungeana,L. davurica and S. davidii. According to the palnt anatomical and physiological characteristicsof environmental adaptation, the testing plants were divided into three types: I, stressavoidance, including A. scoparia, S. bungeana and P. australis; II, strain repair, including A.giraldii, B. ischcemum and L. davurica; III, strain avoidance, A. gmelinii, P. sepium and S.davidii. Integrative subordinative value was calculated based on reproductive, morphological,anatomical and physiological characteristics of plant. The integrative subordinative values oftesting plants on different soil erosion environments of slope-gully system were positivelycorrelated with the species important value in community. From the results of comprehensiveanalysis of the subordinate function, two shrubs, S. davidii and P. sepium, showed higher environmental adaptability than another testing plants, followed by widespread plants (A.gmelinii, L. davurica, S. bungeana), the environmental adaptability of B. ischcemum, A.giraldii, P. australis, A. scoparia which only distributed on certain erosion environments,were relatively weak among testing plants.
（1）由森林带向草原带过渡，植物群落结构变的简单，物种多样性减小，但物种个数反而有增加趋势；中生植物、高位芽植物减少，旱生植物、隐芽植物增加。坡沟系统不同侵蚀环境下物种均匀度差异不显著，但阴坡群落物种多样性和丰富度较阳坡提高；阳沟坡旱生植物和隐芽植物比例大；而峁顶一年生植物比例增大。从阳沟坡至阴沟坡随着侵蚀程度的减小，物种数量和密度增加，但各物种生态位宽度和重要值减小。旱生和旱中生植物在黄土丘陵沟壑区有着广泛的分布。
（2）同种植物单个花序（花）重、单个种子重量表现出一定的遗传稳定性，在坡沟系统不同侵蚀环境下差异不明显。阳沟坡和阴沟坡植物有性生殖投入低，但该侵蚀环境下隐芽植物比例大，使得其上物种主要以芽库完成地上植被更新。峁顶和阳峁坡植物有性生殖投入量大，产生大量的种子有利于植物种群的扩大。相对其他供试物种，达乌里胡枝子有性繁殖分配比例大，而两种灌木狼牙刺与杠柳则相反。在芽库季节动态上，植物返青期较大的永久性芽库存量保证了植物迅速更新，开花结实期大量季节性芽库确保了植物光合产物积累。
（3）同种植物叶片形态解剖结构在3个植被带相同坡沟侵蚀环境下变化差异性不显著，但坡沟系统不同的侵蚀环境下植物叶片形态和解剖结构变化差异明显。随着侵蚀程度的增加，叶片叶片厚度、干物质量积累、角质层厚度、栅栏组织厚度、木质部韧皮部面积比增加，叶面积、表皮细胞大小、海绵组织厚、细胞间隙减小。输送水分的木质部与植物适应环境能力关系密切。不同植物适应黄土高原半干旱的气候也有不同的策略：猪毛蒿叶具有发达贮水组织，铁杆蒿全栅型叶片，茭蒿和狼牙刺具有双栅型叶片和大的维管束鞘细胞，达乌里胡枝子叶具有发达的粘液细胞，白羊草和芦苇叶上表皮具有发达的泡状细胞，长芒草叶上表皮下多层厚壁组织，杠柳厚的角质层。综合隶属函数分析表明，叶片保护组织发达的长芒草从叶片形态与解剖角度分析对环境适应能力最强。
（4）三个植被带相同坡沟侵蚀环境下植物叶片生理特性变化未达到显著性，但坡沟系统不同侵蚀环境下植物叶片生理特性差异明显。从阴沟坡至阳沟坡随着侵蚀程度增加，细胞受伤害和叶绿素受破坏程度增加，但植物通过叶片保水力、渗透调节物质、抗逆酶活性、非酶抗氧化物质的加强来应对环境胁迫，同时减小叶片相对含水量、自由束缚水比值，以牺牲植物生长速率的代价来应对胁迫环境。所测生理指标中，与植物环境适应相关性最大的是抗坏血酸氧化酶。隶属函数综合分析表明两种灌木杠柳和狼牙刺从生理角度分析对环境适应性较强。
（5）不同构型植物拦截坡面径流沉积物和保护植冠下土体而形成不同形态的土堆。随着坡度的增加，土堆由对称堆型圆帽结构转变为不对称的台阶型结构，主杆型狼牙刺转折点在15-25°的陡坡，聚丛型铁杆蒿和簇丛型白羊草在26-35°的极陡坡，疏丛型茭蒿在≥36°的险坡。坡度和植丛顺坡基部直径影响着土堆的高度；而植丛垂直坡面的基部直径、植丛分枝数量及整个植丛的冠幅影响着土堆的面积。随着坡度的增加土堆高度总体上呈增加趋势而土堆面积趋势相反。植物拦截形成的土堆成为坡面的“营养岛”，相对于植被间表现出良好的土壤理化特性及较高的物种多样性。茭蒿以其发达且扩张的枝系、狼牙刺以其较大的植冠在形成土堆和对坡面微环境的改善方面能力较强。
（6）在繁殖策略方面，供试物种猪毛蒿、铁杆蒿、茭蒿、白羊草、杠柳趋向于r-生殖策略，长芒草、达乌里胡枝子、狼牙刺更趋向于K-生殖策略。依据植物适应环境时的形态解剖与生理特点，供试植物可分为Ⅰ：御逆型，猪毛蒿、芦苇、长芒草；Ⅱ：胁变修复型，茭蒿、白羊草、达乌里胡枝子；Ⅲ：御胁变型，铁杆蒿、狼牙刺、杠柳。综合植物繁殖、形态、解剖和生理特征，供试物种综合隶属函数值与物种重要值呈正相关关系，供试植物的侵蚀环境适应能力表现为：两种灌木狼牙刺和杠柳较强，在黄土丘陵沟壑区不同侵蚀环境有着广泛分布的铁杆蒿、达乌里胡枝子、长芒草次之，而对侵蚀环境具有选择性的白羊草、茭蒿、芦苇和猪毛蒿相对较小。
The serious soil erosion on the Loess Plateau cause the deep cut gullies, fragmentation oflandform and form a complicated landform, thus result in spatial heterogeneity of water andheat. Then, there are series of adjustments occur on plant community structure, reproductivestrategies, leaf morphological and physiological characteristics to adapt different soilenvironments. In this study, the community structures in five soil erosion environments onslope-gully system (sunny gully slope, sunny hilly slope, hilltop, shady hilly slope, and shadygully slope) on three vegetation zones (forest zone, forest-steppezone, and steppe zone) on thehilly-gully region of the Loess Plateau were examined. The species niche breadth andimportant value in community were analysed to identify dominant species at different soilerosion environments. The reprodution, morphology, anatomy, and physiology of seclectednine dominant species (Artemisia scoparia, Artemisia gmelinii, Artemisia giraldii,Bothriochloa ischcemum, Stipa bungeana, Phragmites australis, Lespedeza davurica,Periploca sepium and Sophora davidii) were studied. And, four typical shoot architecturespecies (A. gmelinii, with dense shoot architecture; A. giraldii, with expanding shootarchitecture; B. ischaemum, with a tussock-formingshoot architecture; and S. viciifolia, with amain-stem shoot architecture) was selected to analysis the shape of phytogenic mounds andits influence of mirco-environmnent on slope. The main results are as follows:
(1) Fom forest to steppe zone, plant community structures become simple, speciesdiversity decreased, and the number of species showed a slight increased with mesophytesand phanerophytes decreased, while xerophytes and cryptophytes increased. At different soilerosion environments on slope-gully system, the variation of species evenness was notsignificant. On shady slope, the species diversity and richness were relative larger than sunnyslope. The xerophyte and cryptophytes increased at sunny gully slope. Hilltop had a higher therophyte proportion than the other erosion environments. From sunny to shady gully slope,the species niche and important value in community decreased with the decreasing of erosiondegree. Xerophyte and xero-mesophytes have a wide range of distribution on the hilly-gullyregion of the Loess Plateau.
(2) For same species, single dry weight of inflorescence (flower) and seed mass showedcertain genetic stability, and no significant difference existed in different soil erosionenvironments on slope-gully system. At the sunny gully slope and shady gully slope, plantreproductive investment was low, but the high proportion of cryptophytes on theseenvironments made vegetation regeneration relying mainly on bud bank. The reproductiveinvestment of species on hilltop and sunny hilly slope were larger than the other three erosionenvironments which result in a large number of seeds and thus was beneficial to the expansionof seed dispersal plants. L. davurica had a larger reproductive allocation, while two shrubs (P.sepium and S. davidii) were relatively less. In vegetation turning-green season, the largerperennial bud bank ensure a rapid vegetation renewal. The larger seasonal bud bank onfruit-setting season ensure the accumulation of photosynthate.
(3) The leaf morphological and anatomical characteristics at same slope-gully erosionenvironment on the three vegetation zones had a slight changes, but these characteristicsshowed significant changes at different erosion environments on slope-gully system. With theincreasing of soil erosion degree, leaf thickness, dry matter content, cuticle thickness, palisadetissue thickness, xylem area/phloem area increased, while leaf area, epidermal cell size,spongy tissue thickness, intercellular space decreased. In those leaf morphological andanatomical indices, xylem area, which mediate plant water transportation, had the closestrelationship with plant environmental adaptability. Different plants have its special structureto adapt the semi-arid climate on the Loess Plateau: water storage parenchyma of A. scoparia,total-palisade tissue of A. gmelinii, bi-palisade tissue and big bundle sheath cells of A. giraldiiand S. davidii, mucilage cell of L. daurica; special bulliform cells of B. ischcemum and P.australis; multilayer sclerenchyma of S. bungeana; thick cuticle of P. sepium. From leafmorphological and anatomical characteristics, comprehensive analysis of the subordinatefunction showed that S. bungeana, which has a well developed protective tissues, had thestrongest environmental adaptability.
(4) Similar to plant morphological characteristics, plant physiological indices at sameslope-gully erosion environment on the three vegetation zones showed a slight changes, butthose characteristics presented significant changes at different erosion environments on slope-gully system. From sunny to shady gully slope，with the increasing of soil erosiondegree, the degree of cell damage and amount of chlorophyll destruction increased. But plantresistance to environmental stress by increasing leaf water-holding ability, osmotic adjustmentsubstances, stress-resistance enzymes, non-enzymatic antioxidants, meanwhile by decreasingleaf relative water content and free water/bound water. This processes are at the expense ofplant growth rate to deal with the environmental stress. Among the21measured physiologicalindices, ascorbic acid oxidase (APX) had the closest relationship with plant environmentaladaptability. From plant physiological indices, comprehensive analysis of the subordinatefunction showed that two testing shrubs, S. davidii and P. sepium, had a strong environmentaladaptability.
(5) Different shoot architecture species have different ability of preventing soil erosionand retaining sediment, thus the shape of phytogenic mound changes with slope gradientincreasing. Mound shapes changed from symmetrical mound-type structures intoasymmetrical terrace-type structures as slope gradient increased, but the change at0-15°forS. viciifolia, at26-35°for A. gmelinii and B. ischaemum, at>35°for A. giraldii. Among theplant shoot architecture parameters, plant basal diameter along the slope was significantlycorrelated with mound height, while the plant basal diameter perpendicular to the slope andtotal cross-sectional area of the stem at the plant base were associated with mound area.Mound height increased with slope for all species, while mound area showed oppositetendency. The phytogenic mounds can act as ‘resource islands’, thus improved speciesrichness and stress resistance of plant community on slope. A. giraldii with expanding shootarchitecture and S. viciifolia with its large canopy can create relatively large mounds.
(6) In reproductive strategy, K-reproductive strategyare adopted by A. scoparia, A.gmelinii, A.giraldii, B. ischcemum and P. sepium; and r-reproductive strategy by S. bungeana,L. davurica and S. davidii. According to the palnt anatomical and physiological characteristicsof environmental adaptation, the testing plants were divided into three types: I, stressavoidance, including A. scoparia, S. bungeana and P. australis; II, strain repair, including A.giraldii, B. ischcemum and L. davurica; III, strain avoidance, A. gmelinii, P. sepium and S.davidii. Integrative subordinative value was calculated based on reproductive, morphological,anatomical and physiological characteristics of plant. The integrative subordinative values oftesting plants on different soil erosion environments of slope-gully system were positivelycorrelated with the species important value in community. From the results of comprehensiveanalysis of the subordinate function, two shrubs, S. davidii and P. sepium, showed higher environmental adaptability than another testing plants, followed by widespread plants (A.gmelinii, L. davurica, S. bungeana), the environmental adaptability of B. ischcemum, A.giraldii, P. australis, A. scoparia which only distributed on certain erosion environments,were relatively weak among testing plants.
引文
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