三峡库区两种耐水淹植物的存活率和碳水化合物储备关系

野古草和秋花柳是三峡库区消落带两种强水淹耐受能力的植物物种。以往研究显示植物的水淹耐受性和体内碳水化合物储备有关。为了探明野古草和秋花柳水淹下的高存活率是否和碳水化合物储备有关, 研究了在室外6个月的模拟水淹条件下两个物种在不同水淹时间(40、90、120和180d)和不同水淹深度下(不水淹、根部水淹和完全淹没)的生物量积累、存活率和碳水化合物含量和分布。结果表明: (1)野古草和秋花柳对长期水淹具有很高的耐受性, 根部水淹植物6个月处理后完全存活; 而完全淹没条件下, 野古草仅在4个月, 秋花柳仅在6个月处理后才开始死亡; (2)碳水化合物主要储备在野古草的茎和秋花柳的茎与主根中, 野古草的根和秋华柳的细根中碳水化合物含量很低; (3)水淹深度和水淹时间对植物生物量积累和碳水化合物含量影响显著(P 0.05):与未水淹植株相比, 根部水淹仅略微降低了生物量积累以及可溶性糖和淀粉含量 (P 0.05), 且保持基本稳定或增加的趋势, 而完全淹没的植株生物量随水淹时间逐渐降低, 碳水化合物含量在前90天快速下降 (P 0.05), 之后缓慢下降或保持不变。研究结果表明, 野古草和秋花柳强的水淹耐受性是和它们高的碳水化合物储备以及水淹条件下对碳水化合物的动用能力有关, 后期的死亡率增加与碳水化合物储备消耗殆尽有关, 野古草和秋花柳对碳水化合物储备对水淹的响应的差异可能和它们的碳水化合物储备在不同组织中的分配模式有关。       

三峡库区岸生植物野古草(Arundinella anomala Steud.)光合作用对水淹的响应

为了阐明水淹对三峡库区岸生植物野古草光合作用的影响,模拟了三峡库区消落带水淹发生情况,考察了在不同水淹处理下野古草(Arundinella anomalaSteud.)的光合及叶绿素荧光特性。实验设置了对照(不进行水淹,常规供水管理)、半淹(植株置于水中,植株地上部分一半被淹没)、水下0.5m(植株置于水中,植株顶部在水面下0.5m)、水下2m(植株置于水中,植株顶部在水面下2m)4个不同的水淹深度和02、0、40d和60d等4个不同的水淹时间处理,测定了在不同水淹深度和水淹时间处理下野古草的净光合速率、总叶绿素含量、PSⅡ的最大光化学效率、电子传递速率、表观量子效率、叶绿素利用效率与羧化效率。结果发现,在水淹前期,水淹对野古草的光合特性影响较小,直到水淹60d后,才对野古草的光合特性产生明显影响,且影响程度随水淹深度的不同而不同。野古草在水淹20d和水淹40d后,各水淹处理的净光合速率与对照相比无明显降低,其中水淹20d后,半淹处理的野古草叶片净光合速率比对照还高出16.1%。水淹60d后,水下0.5m和水下2m的净光合速率显著低于对照和半淹,其净光合速率分别为7.51μmol.m-2.s-1和9.15μmol.m-2.s-1。结果表明,水淹20d和40d对野古草的电子传递速率、表观量子效率和羧化效率没有影响。水淹处理60d后,与对照植株相比,半淹处理植株的电子传递速率、表观量子效率、叶绿素利用效率和羧化效率没有明显变化,但水下0.5m和水下2m处理植株的电子传递速率、表观量子效率、叶绿素利用效率和羧化效率有明显降低。在整个实验期间,半淹处理植株的净光合速率、电子传递速率、表观量子效率和羧化效率没有受到任何不利影响。尽管在水淹60d后水下0.5m和水下2m处理植株的净光合速率、电子传递速率、表观量子效率、叶绿素利用效率和羧化效率降低,但降低后的数值仍不低于甚至高于一些自然生长的未受水淹的植物物种。研究表明,野古草对水淹具有很好的耐受能力,是一种可以用于三峡库区消落区植被构建的优良植物物种。     

三峡库区岸生植物野古草(Arundinella anomala Steud.)光合作用对水淹的响应

为了阐明水淹对三峡库区岸生植物野古草光合作用的影响,模拟了三峡库区消落带水淹发生情况,考察了在不同水淹处理下野古草(Arundinella anomala Steud.)的光合及叶绿素荧光特性。实验设置了对照 (不进行水淹,常规供水管理)、半淹 (植株置于水中,植株地上部分一半被淹没)、水下0.5m (植株置于水中,植株顶部在水面下0.5m)、水下2m (植株置于水中,植株顶部在水面下2m) 4个不同的水淹深度和0、20、40d和60d等 4个不同的水淹时间处理,测定了在不同水淹深度和水淹时间处理下野古草的净光合速率、总叶绿素含量、PSⅡ的最大光化学效率、电子传递速率、表观量子效率、叶绿素利用效率与羧化效率。结果发现,在水淹前期,水淹对野古草的光合特性影响较小,直到水淹60d后,才对野古草的光合特性产生明显影响,且影响程度随水淹深度的不同而不同。野古草在水淹20d和水淹40d后,各水淹处理的净光合速率与对照相比无明显降低,其中水淹20d后,半淹处理的野古草叶片净光合速率比对照还高出16.1%。水淹60d后,水下0.5m和水下2m的净光合速率显著低于对照和半淹,其净光合速率分别为7.51μmol•m－2•s－1和9.15μmol•m－2•s－1。结果表明,水淹20d和40d对野古草的电子传递速率、表观量子效率和羧化效率没有影响。水淹处理60d后,与对照植株相比,半淹处理植株的电子传递速率、表观量子效率、叶绿素利用效率和羧化效率没有明显变化,但水下0.5m和水下2m处理植株的电子传递速率、表观量子效率、叶绿素利用效率和羧化效率有明显降低。 在整个实验期间,半淹处理植株的净光合速率、电子传递速率、表观量子效率和羧化效率没有受到任何不利影响。尽管在水淹60d后水下0.5m和水下2m处理植株的净光合速率、电子传递速率、表观量子效率、叶绿素利用效率和羧化效率降低,但降低后的数值仍不低于甚至高于一些自然生长的未受水淹的植物物种。研究表明,野古草对水淹具有很好的耐受能力,是一种可以用于三峡库区消落区植被构建的优良植物物种。     

水淹对三峡库区两种岸生植物秋华柳(Salix variegata Franch.)和野古草(Arundinella anomala Steud.)水下光合的影响

为了阐明水淹对三峡库区岸生植物秋华柳(Salix variegata Franch.)和野古草(Arundinella anomala Steud.)水下光合作用的影响,模拟三峡库区消落带水淹发生情况,考察了在不同水淹处理下秋华柳和野古草的水下光合.实验设置了对照(不进行水淹,常规供水管理)和2m深度水淹(植株置于水中,植株顶部在水面下2 m)2个不同的水淹深度和5、15、20、40 d和60 d等5个不同的水淹时间处理,采用Chlorolab-2液相氧电极(英国Hansatech公司生产)测定了在不同水淹时间和水淹深度处理下秋华柳和野古草的水下光合作用.实验结果表明: (1) 水淹60d后,秋华柳和野古草的存活率均为100%,而典型的陆生植物香樟和马唐分别在水淹40d和15d后全部死亡.(2) 在相同的水淹时间和水淹深度下,秋华柳和野古草的水下光合速率(放氧速率)显著高于典型的陆生植物(香樟(Cinnamomum camphora(L.) Presl.)和马唐(Digitaria sanguinalis (L.)Scop.)) (3) 在长期水淹的条件下,秋华柳和野古草仍具有水下光合的能力.在水淹60 d后,水淹2 m的秋华柳和野古草植株的水下放氧速率显著低于对照植株的水下放氧速率,但仍具有水下光合的能力,其水下光合速率分别为0.202 μmol · m-2 · s-1和0.139 μmol · m-2 · s-1.同时,研究也表明在水淹40 d和60 d后,秋华柳表现出比野古草强的水下光合能力.研究表明,秋华柳和野古草在存活率和水下光合方面对长期水淹表现出良好的适应性,是可以用于三峡库区消落区植被构建的优良植物物种.     

长期水淹条件下香根草(Vetiveria zizanioides)、菖蒲(Acorus calamus)和空心莲子草(Alternanthera philoxeroides)的存活及生长响应

人工构建三峡库区消落区植被是控制消落区水土流失、保护消落区生态环境的重要措施,选择能够耐受长时间完全水淹的植物物种是该措施实施的关键.为了验证香根草、菖蒲、空心莲子草能否用于消落区植被的构建,实验模拟消落区的长期完全水淹条件,设置30d、60d、90d、120d、150d和180d等6个完全水淹时间水平,研究了3种植物在完全水淹条件下生长、生物量积累及存活状况.结果发现:(1)3种植物在经受长时间的完全水淹后有较高的存活率,180d全淹处理后,香根草、菖蒲和空心莲子草的存活率分别为87.5%、100%和50%.(2)这3种植物有不同的水下生长能力.全淹条件下,香根草生长缓慢,几乎没有产生新的叶片,总叶长也没有显著变化;菖蒲能够持续产生较对照植株更为细长的叶片,空心莲子草只在水淹初期(30d内)能够快速伸长地上部分的枝条,并迅速产生新叶片,但随水淹时间的延长,总枝条长及总叶片数没有再显著增加.(3)与对照植株相比,全淹处理抑制了3种植物总生物量的增加,但对3种植物的地上、地下部分生物量抑制程度不同.全淹条件下,香根草的地上部分和地下部分生物量与水淹0d水平(水淹处理开始前一天,下同)相比无显著变化,根冠比高于对照植株;菖蒲的地上部分生物量随水淹时间延长而降低,但却高于对照植株,地下部分生物量始终低于水淹0d水平,根冠比低于对照植株;空心莲子草的地上部分生物量与水淹0d水平相比无显著差异,但地下部分生物量与水淹0d水平相比大幅降低,根冠比低于对照植株.结果表明,这3种植物都有很强的水淹耐受能力,可应用于三峡库区消落区植被的构建.同时,发现植物对长期完全水淹的耐受能力很大程度上与植株在水下的生长情况及植株的营养储备水平相关,剧烈的水下生长会消耗大量的营养储备,进而造成植株存活率降低.植株在全淹条件下有限的生长能力及丰富的营养储备可能是耐淹物种的重要特征.     

水淹胁迫诱导喜旱莲子草茎发生类环割效应

在水淹环境中,喜旱莲子草水面上未受淹的茎段常表现出增粗膨大的现象。遭受水淹的植物被淹没的组织和器官会面临氧气缺乏和能量供应不足的问题。植物体内碳水化合物的运输需要消耗能量,当因水淹而使植株被部分淹没（即地下部分全部和地上部分的一部分被淹没）时,由于水淹缺氧导致能量供应不足,碳水化合物在植物被淹组织内的运输可能受限从而在水面上的未淹茎段中积累并对未淹茎段的径向形态产生影响。为探究水淹环境中喜旱莲子草未受淹茎段增粗膨大是否与碳水化合物积累有关,对茎被水淹和茎不受水淹的喜旱莲子草进行对比研究,结果发现：（1）水淹的喜旱莲子草位于水面上的未淹茎段节间平均直径显著大于水面下受淹茎段节间平均直径,未淹茎段与受淹茎段相比发生显著的膨大现象;未水淹的喜旱莲子草其茎的上部茎段节间平均直径与下部茎段节间平均直径相比并无显著差异,上部茎段也无明显膨大现象。（2）水淹的喜旱莲子草位于水面上的未淹茎段碳水化合物含量显著高于未水淹的喜旱莲子草对应茎段的碳水化合物含量。本研究表明,水淹胁迫下喜旱莲子草位于水面上的未受淹茎段中碳水化合物发生积累,导致植株位于水面上的未受淹茎段发生与物理环割后类似的茎膨大现象。     

外源乙烯和醎萘乙酸对三峡库区岸生植物野古草和秋华柳茎通气组织形成的影响

 为了探究乙烯和α-萘乙酸(α-NAA)是否是水淹环境条件下植物体内通气组织形成的直接原因,对三峡库区岸生植物野古草（Arundinella anomala）和秋华柳（Salix variegata） 在无水淹环境条件下施加乙烯利和α-NAA后茎中通气组织的形成情况进行了研究。实验分3种处理：单独用乙烯利溶液处理（浓度分别为0、250和500 mg●L^－1）、单独用α-NAA溶液处理（浓度分别为0、50和100 mg●L^－1）和二者混合处理（250 mg●L^－1乙烯利溶液 +50 mg●L^－1 α-NAA溶液）。处理5 d后,采用切片法制备其茎中部横切面切片,用E80i Nikon显微镜进行观察,并运用ACT-2U和Simple PCI软件分析野古草和秋华柳茎中通气组织的形成情况。结果显示：在这3种处理条件下,野古草和秋华柳茎中通气组织形成均有明显增强,并且较高浓度的乙烯利溶液促使茎通气组织形成更多,施加的α-NAA浓度越高,形成通气组织的能力越强;混合溶液处理与单独施加250 mg●L^－1乙烯利或单独施加50 mg●L^－1 NAA的处理相比,对通气组织形成的增强效应无明显差异。研究表明,在水淹条件下植物体内通气组织的发生与乙烯和生长素含量的增加有直接关系。     

外源乙烯和醎萘乙酸对三峡库区岸生植物野古草和秋华柳茎通气组织形成的影响

为了探究乙烯和α-萘乙酸(α-NAA)是否是水淹环境条件下植物体内通气组织形成的直接原因,对三峡库区岸生植物野古草（Arundinella anomala）和秋华柳（Salix variegata） 在无水淹环境条件下施加乙烯利和α-NAA后茎中通气组织的形成情况进行了研究。实验分3种处理：单独用乙烯利溶液处理（浓度分别为0、250和500 mg●L^－1）、单独用α-NAA溶液处理（浓度分别为0、50和100 mg●L^－1）和二者混合处理（250 mg●L^－1乙烯利溶液 +50 mg●L^－1 α-NAA溶液）。处理5 d后,采用切片法制备其茎中部横切面切片,用E80i Nikon显微镜进行观察,并运用ACT-2U和Simple PCI软件分析野古草和秋华柳茎中通气组织的形成情况。结果显示：在这3种处理条件下,野古草和秋华柳茎中通气组织形成均有明显增强,并且较高浓度的乙烯利溶液促使茎通气组织形成更多,施加的α-NAA浓度越高,形成通气组织的能力越强;混合溶液处理与单独施加250 mg●L^－1乙烯利或单独施加50 mg●L^－1 NAA的处理相比,对通气组织形成的增强效应无明显差异。研究表明,在水淹条件下植物体内通气组织的发生与乙烯和生长素含量的增加有直接关系。     

水淹对三峡库区岸生植物秋华柳(Salix variegata Franch.)存活和恢复生长的影响

植物在水淹后的存活和恢复生长状况可以衡量其对水淹的耐受能力,为了解三峡库区岸生植物秋华柳(Salix variegata Franch.)的水淹耐受能力,研究了长期水淹条件下秋华柳植株的存活和恢复生长状况.实验设置了3个水淹深度:对照,水淹根部(植株置于水中,仅地下部分被淹没)和水淹2m(植株置于水中,顶部距水面2m),6个水淹时间:20d,40d,60d,90d,120d和180d.研究结果表明:(1)水淹对秋华柳植株存活率的影响较小.水淹根部处理的植株在水淹180d后,存活率仍为100%,水淹2m处理的植株在水淹120d后,存活率也为100%,直到180d后,存活率才下降为0.(2)水淹后,秋华柳植株仍然可以进行恢复生长,表现出很强的恢复生长能力,但因水淹处理的不同,其恢复生长存在差异.随着水淹时间的延长,秋华柳植株出水后到开始恢复生长之前所需的时间增加,但所有水淹处理的植株在水淹结束后一周内都可以开始恢复生长.在相同水淹时间处理下,水淹处理的秋华柳植株在恢复生长期间的相对生长速率都高于对照植株,水淹40d,60d,90d后,水淹2m的秋华柳植株分别比对照植株高57.8%,143.4%,130.4%.水淹结束时,秋华柳地上部分生物量随水淹深度的不同而不同,水淹根部处理的植株几乎与对照植株无显著差异,水淹2m处理的植株都低于对照植株.水淹结束后,不同处理的秋华柳植株生长2个月后的地上部分生物量与其在水淹结束时不同处理植株地上部分生物量的变化趋势相似.本研究表明,秋华柳在长时间的水淹后具有很高的存活率,并可以进行很好的恢复生长,表现出较强的水淹耐受能力,可以考虑将其应用于三峡库区消落区的植被构建.     

EFFECTS OF LONG-TERM SUBMERGENCE ON SURVIVAL AND RECOVERY GROWTH OF FOUR RIPARIAN PLANT SPECIES IN THREE GORGES RESERVOIR REGION, CHINA

 地瓜藤(Ficus tikoua)、荻(Triarrhena sacchariflora)、牛鞭草(Hemarthria altissima)和狗牙根(Cynodon dactylon)是三峡库区常见的岸生植物, 自然分布于河岸带不同垂直高程的地段。为了明确它们在成库后“三峡库区消落区”长期完全水淹条件下的存活和生长情况, 实验设置对照(不进行水淹, 常规供水管理)和完全水淹两个处理, 30、60、90、120、150和180 d 6个水淹时间水平, 研究了4种植物在完全水淹条件下的存活、生物量变化和恢复生长。结果发现: 1) 4种植物在完全水淹条件下的存活率与其在河岸带上的垂直分布高程密切相关。分布于距江面高程较高的河岸段的地瓜藤植株, 在全淹30 d后就全部死亡; 分布在中高程河岸段的荻在全淹150和180 d后全部死亡; 可以分布于低高程河岸段的牛鞭草和狗牙根, 淹没180 d后存活率分别为90%和100%。2)全淹抑制了荻、牛鞭草和狗牙根的生长, 总生物量增量显著低于对照植株。与水淹0 d相比, 全淹处理植株的地上部分生物量显著降低, 荻在全淹60和120 d后, 地下部分生物量显著降低, 但牛鞭草和狗牙根的地下部分生物量与水淹0 d水平相比无显著差异。3)水淹处理结束后, 存活的荻、牛鞭草和狗牙根植株都能很好地恢复生长。在恢复生长过程中, 全淹30、60和90 d后, 荻、牛鞭草和狗牙根植株的总分枝长相对生长速率与对照植株无显著差异, 全淹120、150和180 d后, 牛鞭草和狗牙根植株的总分枝长相对生长速率显著高于对照植株。全淹处理的荻、牛鞭草和狗牙根植株的总叶片数相对生长速率始终显著高于对照植株。遭受长期完全水淹后, 植株在有限的营养储备条件下, 快速产生叶片以迅速积聚光合产物可能是植物更为优化的恢复生长方式。     

Adaptive strategies to tolerate prolonged flooding in seedlings of floodplain and upland populations of Himatanthus sucuuba, a Central Amazon tree

Long-term flooding imposes a strong selection pressure on plants for the development of protective mechanisms to alleviate the harmful effects of hypoxic and anoxic conditions. This is particularly critical in the Amazonian floodplains where plants withstand annual periods of flooding lasting 7 months and mean flooding amplitude reaching 10 m or more. Himatanthus sucuuba (Apocynaceae) is a tree that is found in the varzea (VZ) floodplains and non-flooded terra firme (TF) forests. It was examined whether individuals from these two contrasting habitats respond differently when subjected to extreme flooding conditions. TF and VZ seedlings were experimentally well-watered, waterlogged (roots and parts of the stems flooded), or submerged (whole plant flooded) during a 4-month period. Anaerobic respiration, evaluated by measuring alcohol dehydrogenase (ADH) activity, and root carbohydrate reserves were quantified, given that the availability of readily fermentable carbohydrates is essential to sustain an active fermentative metabolism. We also assessed changes in morphoanatomy, seedling survival, biomass accumulation and distribution. VZ seedlings had greater root concentrations of soluble sugars and starch, larger seedling mass and accumulated more biomass in roots and stems while TF seedlings allocated more towards stem and leaves. ADH activity was low in seedlings of both populations before exposure to flooding. Waterlogging induced an increase in ADH activity that reached a maximum value in 15 days. Thereafter activity decreased slowly, meanwhile a rapid formation of lenticels, adventitious roots and aerenchyma was observed. Submergence induced leaf shedding and the development of aerenchyma in the root cortex. While VZ seedlings maintained high levels of ADH activity throughout the whole 4-month period, ADH activity in TF seedlings peaked about 15 days after submersion followed by a continuous decrease and death of all the plants. Thus, VZ and TF seedlings differed considerably in terms of tolerating long-term exposure to flooding, especially under total submersion. These results suggest that the predictability and long-term duration of flooding in Central Amazon rivers can impose a selective pressure that is strong enough to result in large phenotypic differences between the two populations of H. sucuuba in the two habitat types.     

Formation and function of secondary aerenchyma in hypocotyl, roots and nodules of soybean (Glycine max) under flooded conditions

Flooding is a major problem in many areas of the world and soybean is susceptible to the stress. Understanding the morphological mechanisms of flooding tolerance is important for developing flood-tolerant genotypes. We investigated secondary aerenchyma formation and function in soybean (Glycine max) seedlings grown under flooded conditions. Secondary aerenchyma, a white and spongy tissue, was formed in the hypocotyl, tap root, adventitious roots and root nodules after 3 weeks of flooding. Under irrigated conditions aerenchyma development was either absent or rare and phellem was formed in the hypocotyl, tap root, adventitious roots and root nodules. Secondary meristem partially appeared at the outer parts of the interfascicular cambium and girdled the stele, and then cells differentiated to construct secondary aerenchyma in the flooded hypocotyl. These morphological changes proceeded for 4 days after the initiation of the flooding. After 14 days of treatment, porosity exceeded 30% in flooded hypocotyl with well-developed secondary aerenchyma, while it was below 10% in hypocotyl of irrigated plants that had no aerenchyma. When Vaseline was applied to the hypocotyl of plants from a flooded treatment to prevent the entry of atmospheric oxygen into secondary aerenchyma, plant growth, especially that of roots, was sharply inhibited. Thus secondary aerenchyma might be an adaptive response to flooding.     

Survival of Ranunculus repens L. (creeping buttercup) in an amphibious habitat

The turlough form of Ranunculus repens is subjected to several months' complete inundation with hard groundwater. Experimental flooding to the level of the soil surface had no effect on turlough or ruderal populations relative to drained controls. Experimental submergence resulted in direct tissue death of the ruderal population but did not affect the turlough population relative to drained controls. There was no detectable difference in the proportion of aerenchyma in drained, flooded and submerged roots of plants from either population. The proportion of aerenchyma increased with root age in the ruderal population. Up to twice the proportion of aerenchyma occurred in the lower third of the root in the turlough population relative to the middle and upper thirds. Submergence in artificially hardened tap water increased the amount of tissue death in the ruderal population, whereas it appeared to enhance the growth of plants from the turlough population relative to that of plants submerged in tap water. Only the ruderal population demonstrated a depth accommodation response in submerged conditions. Root concentrations of ethanol-soluble carbohydrates were up to three times higher in a field- collected turlough population during winter and autumn months than those in a ruderal population. Low levels of ethanol-insoluble carbohydrates were present in the turlough population but were absent from the ruderal population. Starch concentrations fluctuated greatly in the turlough population and were generally higher than those in the ruderal population. These results, together with those from previous investigations, suggest that the turlough population survives prolonged submergence by maintaining low levels of submerged photosynthesis, which may circulate oxygen within the plant tissues, and by utilizing storage carbohydrates for maintenance respiration.     

Morphological and photosynthetic responses of riparian plant Distylium chinense seedlings to simulated Autumn and Winter flooding in Three Gorges Reservoir Region of the Yangtze River, China

To evaluate the tolerance of riparian plant Distylium chinense in Three Gorges Reservoir Region to anti-season flooding, a simulation flooding experiment was conducted during Autumn and Winter, and morphology and photosynthesis of D. chinense seedlings and their recovery growth after soil drainage were analyzed in different duration of flooding and flooding depth. The seedlings were submitted to four treatments: (1) 40 seedlings unflooded and watered daily as control (Unflooded, CK); (2) 120 seedlings flooded at 1 cm above the ground level (F-1 cm); (3) 120 seedlings flooded at 12 cm above the ground level (F-12 cm) and (4) 120 seedlings completely submerged with 2 m water depth (F-2 m, top of plants at 2 m below water surface). The flooding survival, plant height, stem diameter, adventitious roots, stem lenticels, epicormic shoots, chlorophyll content and photosynthesis parameters were determined at 0, 15, 30, 90 days in flooding stress and 15, 60 days after soil drainage. The results showed that the survival of the seedlings subjected to flooding was 100% for all repeated measurements in all treatments. Adventitious roots, hypertrophied lenticels and stem hypertrophy were observed in the seedlings flooded for more than 15 d, and increased with the prolonged flooding duration, while disappeared after the soil was drained. Flooding duration and flooding depth showed significant individual and interactive effects on leaf chlorophyll a (Chl a), chlorophyll b (Chl b), and their ratio, chlorophyll (a + b), the net photosynthesis rate (P_n), transpiration rate (T_r), stomatal conductance (C_s), and inter-cellular CO₂ concentration (C_i) of D. chinense seedlings (P < 0.01). After 15 days of flooding, there was no significant decrease in P_n of the flooded seedlings as compared with that of the control seedlings. P_n of the flooded seedlings was significantly lower than that of the control seedlings after 30 days of flooding (P < 0.05), whereas P_n showed no significant difference among seedlings from three flooding depths. After 90 days of flooding, P_n of the F-2 m flooded seedlings was significantly lower than that of the controls, F-1 cm and F-12 cm flooded seedlings (P < 0.05), but still maintained high photosynthetic capacity. P_n of the F-1 cm and F-12 cm flooded seedlings rose gradually after soil drainage, while, it was significantly lower than that of the control seedlings after 15 days of recovery (P < 0.05). After 60 days of recovery, P_n of all seedlings flooded with different depths showed no significant difference as compared with that of the control seedlings and new leaves grew out in the F-2 m flooded seedlings. The effect of all flooding treatments on Gs, T_r, Chl a, Chl b, Chl a/Chl b and chl (a + b) was basically the same as their effect on P_n, while the effect of all flooding treatments on C_i was quite the contrary. Correlation analysis showed that P_n was positively relative with Gs, T_r, Chl a, Chl b and chl (a + b) (P < 0.05) and significantly negative with C_i (P < 0.05). Therefore, the present study demonstrates that D. chinense has high survival and good recovery growth after long-term flooding in anti-season flooding and could be taken as an excellent candidate species in the re-vegetation of water-level-fluctuation areas in Three Gorges Reservoir Region.     

Partial flooding enhances aeration in adventitious roots of black willow (Salix nigra) cuttings

Black willow (Salix nigra) cuttings are used for streambank stabilization where they are subjected to a range of soil moisture conditions including flooding. Flooding has been shown to adversely impact cutting performance, and improved understanding of natural adaptations to flooding might suggest handling and planting techniques to enhance success. However, data assessing the root aeration in adventitious roots that are developed on cuttings of woody species are scant. In addition, it appears that no data are available regarding aeration of the root system under partially flooded conditions. This experiment was designed to examine the effects of continuous flooding (CF) and partial flooding (PF) on aerenchyma formation and radial oxygen loss (ROL) in black willow cuttings. Photosynthetic and growth responses to these conditions were also investigated. Under laboratory condition, replicated potted cuttings were subjected to three treatments: no flooding (control, C), CF, and PF. Water was maintained above the soil surface in CF and at 10 cm depth in PF. Results indicated that after the 28-d treatments, root porosity ranged between 28.6% and 33.0% for the CF and C plants but was greater for the PF plants (39.2% for the drained and 37.2% for the flooded portions). A similar response pattern was found for ROL. In addition, CF treatment led to decreases in final root biomass and root/shoot ratio. Neither CF nor PF had any detectable adverse effects on plant gas exchange or photosystem II functioning. Our results indicated that S. nigra cuttings exhibited avoidance mechanisms in response to flooding, especially the partially flooded condition which is the most common occurrence in riparian systems.     

Internal aeration and respiration of submerged tomato hypocotyls are enhanced by ethylene-mediated aerenchyma formation and hypertrophy

With the impending threat that climate change is imposing on all terrestrial ecosystems, the ability of plants to adjust to changing environments is, more than ever, a very desirable trait. Tomato (Solanum lycopersicum L.) plants display a number of responses that allow them to survive under different abiotic stresses such as flooding. We focused on understanding the mechanism that facilitates oxygen diffusion to submerged tissues and the impact it has on sustaining respiration levels. We observed that, as flooding stress progresses, stems increase their diameter and internal porosity. Ethylene triggers stem hypertrophy by inducing cell wall loosening genes, and aerenchyma formation seems to involve programmed cell death mediated by hydrogen peroxide. We finally assessed whether these changes in stem morphology and anatomy are indeed effective to restore oxygen levels in submerged organs. We found that aerenchyma formation and hypertrophy not only increase oxygen diffusion toward the base of the plant, but also result in an augmented respiration rate. We consider that this response is crucial to maintain adventitious root development under such conditions and, therefore, making it possible for the plant to survive when the original roots die.     

Aerenchyma formation and recovery from hypoxia of the flooded root system of nodulated soybean

BACKGROUND AND AIMS: Flooding results in hypoxia of the root system to which N2 fixation of nodulated roots can be especially sensitive. Morphological adaptions, such as aerenchyma formation, can facilitate the diffusion of oxygen to the hypoxic tissues. Using soybean, the aim of the study was to characterize the morphological response of the nodulated root system to flooding and obtain evidence for the recovery of N metabolism. METHODS: Sections from submerged tissues were observed by light microscopy, while sap bleeding from the xylem was analysed for nitrogenous components. KEY RESULTS: Flooding resulted in the rapid formation of adventitious roots and aerenchyma between the stem (immediately above the water line), roots and nodules. In the submerged stem, taproot, lateral roots and adventitious roots, lysigenous aerenchyma arose initially in the cortex and was gradually substituted by secondary aerenchyma arising from cells derived from the pericycle. Nodules developed aerenchyma from cells originating in the phellogen but nodules situated at depths greater than 7-8 cm showed little or no aerenchyma formation. As a result of aerenchyma formation, porosity of the taproot increased substantially between the 4th and 7th days of flooding, coinciding with the recovery of certain nitrogenous products of N metabolism of roots and nodules transported in the xylem. Thus, on the first day of flooding there was a sharp decline in xylem ureides and glutamine (products of N2 fixation), together with a sharp rise in alanine (product of anaerobic metabolism). Between days 7 and 10, recovery of ureides and glutamine to near initial levels was recorded while recovery of alanine was partial. CONCLUSIONS: N metabolism of the nodulated soybean root system can recover at least partially during a prolonged period of flooding, a process associated with aerenchyma formation.     

Ethylene Accumulation in Flooded Plants

Ethylene concentration in sunflower (Helianthus annuus L.) cuttings increased 5-fold within 6 h after submersion in distilled water and then declined. When only the basal half of the cutting was steeped in water, ethylene concentration was slightly over half the concentration of the completely submerged cutting. Ethylene concentration also increased when cuttings were wrapped with moist paper tissue. When wrapped with Saran transparent plastic film, ethylene concentration increased continuously for 12 h. When part of the stem of an intact plant was wrapped with Saran, ethylene also increased in that part of the stem. When wrapping was removed or submersion was discontinued, accumulated ethylene in the cuttings decreased, much faster from unwrapped cuttings than from previously submerged ones. During 3 h submersion, ethylene production rate in submerged cuttings was approximately 10% of that for the controls and over 97% ethylene escaped out of the control cuttings while only 22-52% escaped from the submerged cuttings. Water content increased during submersion and decreased when submersion was discontinued. Water content did not change significantly during wrapping, but decreased when the cuttings were unwrapped. High water content in the submerged cuttings was apparently not related to the high ethylene concentration in the cuttings. Causes of ethylene increase in flooded plants were discussed and it was concluded that one of the first and major causes is the accumulation of ethylene in flooded portions of the plants due to the blockade of ethylene escape by water.