Allometric growth, disturbance regime, and dilemmas of controlling invasive plants: a model analysis

Disturbed communities are observed to be more susceptible to invasion by exotic species, suggesting that some attributes of the invaders may interact with disturbance regime to facilitate invasion success. Alternanthera philoxeroides, endemic to South America, is an amphibious clonal weed invading worldwide. It tends to colonize disturbed habitats such as riparian zones, floodplain wetlands and agricultural areas. We developed an analytical model to explore the interactive effects of two types of physical disturbances, shoot mowing and root fragmentation, on biomass production dynamics of A. philoxeroides. The model is based on two major biological assumptions: (1) allometric growth of root (belowground) vs. shoot (aboveground) biomass and (2) exponential regrowth of shoot biomass after mowing. The model analysis revealed that the interaction among allometric growth pattern, shoot mowing frequency and root fragmentation intensity might lead to diverse plant ‘fates’. For A. philoxeroides whose root allocation decreases with growing plant size, control by shoot mowing was faced with two dilemmas. (1) Shoot regrowth can be effectively suppressed by frequent mowing. However, frequent shoot mowing led to higher biomass allocation to thick storage roots, which enhanced the potential for faster future plant growth. (2) In the context of periodic shoot mowing, individual shoot biomass converged to a stable equilibrium value which was independent of the root fragmentation intensity. However, root fragmentation resulted in higher equilibrium population shoot biomass and higher frequency of shoot mowing required for effective control. In conclusion, the interaction between allometric growth and physical disturbances may partially account for the successful invasion of A. philoxeroides; improper mechanical control practices could function as disturbances and result in exacerbated invasion.     

Specialist Insect Herbivore and Light Availability Do Not Interact in the Evolution of an Invasive Plant

Release from specialist insect herbivores may allow invasive plants to evolve traits associated with decreased resistance and increased competitive ability. Given that there may be genetic trade-off between resistance and tolerance, invasive plants could also become more tolerant to herbivores. Although it is widely acknowledged that light availability affects tolerance to herbivores, little information is available for whether the effect of light availability on tolerance differ between the introduced and native populations. We conducted a common garden experiment in the introduced range of Alternanthera philoxeroides using ten invasive US and ten native Argentinean populations at two levels of light availability and in the presence or absence of a specialist stem-boring insect Agasicles hygrophila. Plant biomass (total and storage root biomass), two allocation traits (root/shoot ratio and branch intensity, branches biomass/main stem biomass) and two functional traits (specific stem length and specific leaf area), which are potentially associated with herbivore resistance and light capture, were measured. Overall, we found that A. philoxeroides from introduced ranges had comparable biomass and tolerance to specialist herbivores, lower branch intensity, lower specific stem length and specific leaf area. Moreover, introduced populations displayed higher shade tolerance of storage root biomass and lower plastic response to shading in specific stem length. Finally, light availability had no significant effect on evolution of tolerance to specialist herbivores of A. philoxeroides. Our results suggest that post-introduction evolution might have occurred in A. philoxeroides. While light availability did not influence the evolution of tolerance to specialist herbivores, increased shade tolerance and release from specialist insects might have contributed to the successful invasion of A. philoxeroides.     

水体溶氧影响陆生植物喜旱莲子草(Alternanthera philoxeroides)和牛鞭草(Hemarthria altissima)对完全水淹的耐受力

溶氧是水环境中一个重要的环境因子,为了探讨水中的溶氧含量水平是否会对陆生植物的耐淹能力造成影响,研究了陆生植物喜旱莲子草(Alternanthera philoxeroides)和牛鞭草(Hemarthria altissima)在遭受不同溶氧含量水体完全淹没后的生长表现、存活情况和非结构碳水化合物的变化。实验结果表明:(1)水体中的溶氧含量显著影响了处于完全水淹环境中的喜旱莲子草和牛鞭草的存活。受高溶氧水体完全水淹的喜旱莲子草和牛鞭草主茎的完好程度和存活叶的数量均显著高于遭受低溶氧水体完全水淹的喜旱莲子草和牛鞭草,喜旱莲子草和牛鞭草在高溶氧水体完全水淹后的生物量比低溶氧水体完全水淹后要高;(2)水体中的溶氧含量显著影响了处于完全水淹环境中的喜旱莲子草和牛鞭草的生长,受高溶氧水体完全水淹的喜旱莲子草主茎伸长生长和不定根生长显著强于受低溶氧水体完全水淹的喜旱莲子草,在不定根的生长上牛鞭草也具有同样的表现。(3)高溶氧水环境有利于减小被完全淹没的喜旱莲子草和牛鞭草的碳水化合物消耗,两种植物在受高溶氧完全水淹后体内具有的非结构性碳水化合物含量均比受低溶氧完全水淹后高。(4)喜旱莲子草比牛鞭草能更好地耐受完全水淹,当处于低溶氧完全水淹时表现得更为明显,本研究表明入侵物种喜旱莲子草比本地物种牛鞭草具有更强的环境适应能力和水淹耐受能力。     

Shifting effects of physiological integration on performance of a clonal plant during submergence and de-submergence

Background and Aims

Submergence and de-submergence are common phenomena encountered by riparian plants due to water level fluctuations, but little is known about the role of physiological integration in clonal plants (resource sharing between interconnected ramets) in their adaptation to such events. Using Alternanthera philoxeroides (alligator weed) as an example, this study tested the hypotheses that physiological integration will improve growth and photosynthetic capacity of submerged ramets during submergence and will promote their recovery following de-submergence.

Methods

Connected clones of A. philoxeroides, each consisting of two ramet systems and a stolon internode connecting them, were grown under control (both ramet systems untreated), half-submerged (one ramet system submerged and the other not submerged), fully submerged (both ramet systems submerged), half-shaded (one ramet system shaded and the other not shaded) and full-shaded (both ramet systems shaded) conditions for 30 d and then de-submerged/de-shaded for 20 d. The submerged plants were also shaded to very low light intensities, mimicking typical conditions in turbid floodwater.

Key Results

After 30 d of submergence, connections between submerged and non-submerged ramets significantly increased growth and carbohydrate accumulation of the submerged ramets, but decreased the growth of the non-submerged ramets. After 20 d of de-submergence, connections did not significantly affect the growth of either de-submerged or non-submerged ramets, but de-submerged ramets had high soluble sugar concentrations, suggesting high metabolic activities. The shift from significant effects of integration on both submerged and non-submerged ramets during the submergence period to little effect during the de-submergence period was due to the quick recovery of growth and photosynthesis. The effects of physiological integration were not found to be any stronger under submergence/de-submergence than under shading/de-shading.

Conclusions

The results indicate that it is not just the beneficial effects of physiological integration that are crucial to the survival of riparian clonal plants during periods of submergence, but also the ability to recover growth and photosynthesis rapidly after de-submergence, which thus allows them to spread.     

Recovery dynamics of growth, photosynthesis and carbohydrate accumulation after de-submergence: a comparison between two wetland plants showing escape and quiescence strategies

Background and Aims

The capacity for fast-growth recovery after de-submergence is important for establishment of riparian species in a water-level-fluctuation zone. Recovery patterns of two wetland plants, Alternanthera philoxeroides and Hemarthria altissima, showing ‘escape’ and ‘quiescence’ responses, respectively, during submergence were investigated.

Methods

Leaf and root growth and photosynthesis were monitored continuously during 10 d of recovery following 20 d of complete submergence. Above- and below-ground dry weights, as well as carbohydrate concentrations, were measured several times during the experiment.

Key Results

Both species remobilized stored carbohydrate during submergence. Although enhanced internode elongation depleted the carbohydrate storage in A. philoxeroides during submergence, this species resumed leaf growth 3 d after de-submergence concomitant with restoration of the maximal photosynthetic capacity. In contrast, some sucrose was conserved in shoots of H. altissima during submergence, which promoted rapid re-growth of leaves 2 d after de-submergence and earlier than the full recovery of photosynthesis. The recovery of root growth was delayed by 1–2 d compared with leaves in both species.

Conclusions

Submergence tolerance of the escape and quiescence strategies entails not only the corresponding regulation of growth, carbohydrate catabolism and energy metabolism during submergence but also co-ordinated recovery of photosynthesis, growth and carbohydrate partitioning following de-submergence.     

Spatial arrangements affect suppression of invasive Alternanthera philoxeroides by native Hemarthria compressa

It has been hypothesized that differences in spatial arrangements change the relative frequency of intra- and interspecific encounters between plant species. Manipulating spatial arrangement may play a role in invasive plant suppression when native species are used as competitors against introduced species. In this study, a replacement series experiment was performed to investigate the effects of intraspecifically random and aggregated spatial arrangements on interactions between the native plant Hemarthria compressa and the invasive plant Alternanthera philoxeroides, to test the possibility and effectiveness of H. compressa in suppressing A. philoxeroides. When both species were planted in intraspecifically random spatial patterns, H. compressa had a competitive advantage over A. philoxeroides at relative densities of 2:2 and 3:1. However, aggregation increased the strength, and therefore the cost, of intraspecific competition in H. compressa, resulting in lower biomass production, which reduced its effectiveness as an interspecific competitor. As the relative density of H. compressa in mixtures decreased, plants allocated more biomass to belowground parts, but fewer interspecific encounters lowered its inhibitory effects on A. philoxeroides. The results not only confirm that the frequency of conspecific and heterospecific encounters can influence competitive outcomes, but also suggest that a reduction in the degree of spatial aggregation in H. compressa and an increase in its relative densities may be essential to increase the suppression of A. philoxeroides.     

Phenotypic variation and water selection potential in the stem structure of invasive alligator weed

The morphological and anatomical characteristics of stems have been found to be related to drought resistance in plants. Testing the phenotypic selection of water availability on stem anatomical traits would be useful for exploring the evolutionary potential of the stem in response to water availability. To test the phenotypic variation of the stem anatomical traits of an invasive plant in response to water availability, we collected a total of 320 individuals of Alternanthera philoxeroides from 16 populations from terrestrial and aquatic habitats in 8 plots in China and then analyzed the variation, differentiation, plasticity and selection potential of water availability on the stem anatomical traits. We found that except for the thickness of the cortex, all of the examined phenotypic parameters of the A. philoxeroides stem were significantly and positively correlated with soil water availability. The phenotypic differentiation coefficient for all of the anatomical structural parameters indicated that most of the variation existed between habitats within the same plot, whereas there was little variation among plots or among individuals within the same habitat except for variation in the thickness of the cortex. A significant phenotypic plasticity response to water availability was found for all of the anatomical traits of A. philoxeroides stem except for the thickness of the cortex. The associations between fitness and some of the anatomical traits, such as the stem diameter, the cortex area-to-stem area ratio, the pith cavity area-to-stem area ratio and the density of vascular bundles, differed with heterogeneous water availability. In both the aquatic and terrestrial habitats, no significant directional selection gradient was found for the stem diameter, the cortex area-to-stem area ratio or the density of vascular bundles. These results indicated that the anatomical structure of the A. philoxeroides stem may play an important role in the adaptation to changes in water availability.     

Morphological changes and resource allocation of Zizania latifolia (Griseb.) Stapf in response to different submergence depth and duration

Plants around ponds, rivers and lakes are subjected to long-term partial or complete submergence. When they are flooded, water level affects the plants simultaneously with duration of submergence. Separate and interactive effects of water level and duration on the growth of the herbaceous perennial Zizania latifolia (Poaceae) were investigated by exposing the plants in greenhouse water tanks to submergence in different water depths and for different time-spans. The plants exhibited great shoot elongation upon submergence and prolonged flood duration, and the basal tiller number of the species decreased with higher water levels. Submergence treatment advanced the flowering date and increased the inflorescence number. Plant total biomass did not differ among all the treatments, while the root:shoot ratio decreased with increased water level, prolonged duration of submergence and their interaction. The high plasticity in morphology and shifts in reproductive strategy and biomass allocation enabled the Zizania plants to survive the compound effect of flooding height and duration. This may explain the occurrence of this species in habitats subjected to long-term flooding. The results obtained in this experiment will contribute to understanding the impact of flooding dynamics on plants and the ways of adaptation responses to prolonged waterlogging.     

外源钙离子对南方菟丝子寄生入侵植物喜旱莲子草效果的影响

外源钙离子可以有效地缓解植物在胁迫下受到的毒害,提高植物组织或细胞抗胁迫的能力。以喜旱莲子草为研究对象,采用完全随机区组的实验设计,研究不同钙离子浓度与南方菟丝子寄生对喜旱莲子草茎生物量的影响,探讨外源钙离子是否可以减少南方菟丝子对喜旱莲子草的损伤。研究表明南方菟丝子寄生对喜旱莲子草的根、茎和叶的生物量具有显著影响,且南方菟丝子寄生与外源钙离子对喜旱莲子草茎和叶的生长具有显著的交互作用。不同浓度的钙离子与南方菟丝子寄生对喜旱莲子草茎生物量存在拮抗的交互作用;而仅8 mmol/L Ca2+与南方菟丝子寄生对喜旱莲子草叶生物量存在拮抗的交互作用。不同浓度钙离子均可以缓减南方菟丝子对喜旱莲子草茎的有害效应,仅8 mmol/L Ca2+可以缓减南方菟丝子对喜旱莲子草根和叶的有害效应。2—6 mmol/L的外源钙离子对南方菟丝子寄生下的喜旱莲子草的丙二醛(MDA)含量无显著影响,但可以显著地降低过氧化物酶(POD)活性,表明低浓度的外源钙离子的添加可以缓减南方菟丝子对喜旱莲子草的寄生胁迫,并体现在生理指标上。     

The Invasive Plant Alternanthera philoxeroides Was Suppressed More Intensively than Its Native Congener by a Native Generalist: Implications for the Biotic Resistance Hypothesis

Prior studies on preferences of native herbivores for native or exotic plants have tested both the enemy release hypothesis and the biotic resistance hypothesis and have reported inconsistent results. The different levels of resistance of native and exotic plants to native herbivores could resolve this controversy, but little attention has been paid to this issue. In this study, we investigated population performance, photosynthesis, leaf nitrogen concentration, and the constitutive and induced resistances of the successful invasive plant, Alternanthera philoxeroides, and its native congener, Alternanthera sessilis, in the presence of three population densities of the grasshopper, Atractomorpha sinensis. When the grasshopper was absent, leaf biomass, total biomass, photosynthesis, and leaf nitrogen concentration of A. philoxeroides were higher than those of A. sessilis. However, the morphological and physiological performances of A. philoxeroides were all decreased more intensively than A. sessilis after herbivory by grasshoppers. Especially as the concentrations of constitutive lignin and cellulose in leaf of A. philoxeroides were higher than A. sessilis, A. philoxeroides exhibited increased leaf lignin concentration to reduce its palatability only at severe herbivore load, whereas, leaf lignin, cellulose, and polyphenolic concentrations of A. sessilis all increased with increasing herbivory pressure, and cellulose and polyphenolic concentrations were higher in A. sessilis than in A. philoxeroides after herbivory. Our study indicated that the capability of the invasive plant to respond to native insect damage was lower than the native plant, and the invasive plant was suppressed more intensively than its native congener by the native insect. Our results support the biotic resistance hypothesis and suggest that native herbivores can constrain the abundance and reduce the adverse effects of invasive species.     

Growth promotion and inhibition of the Amazonian wild rice species Oryza grandiglumis to survive flooding

In Asian cultivated rice (Oryza sativa), distinct mechanisms to survive flooding are activated in two groups of varieties. Submergence-tolerant rice varieties possessing the SUBMERGENCE1A (SUB1A) gene display reduced growth during flash floods at the seedling stage and resume growth after the flood recedes, whereas deepwater rice varieties possessing the SNORKEL1 (SK1) and SNORKEL2 (SK2) genes display enhanced growth based on internodal elongation during prolonged submergence at the mature stage. In this study, we investigated the occurrence of these growth responses to submergence in the wild rice species Oryza grandiglumis, which is native to the Amazon floodplains. When subjected to gradual submergence, adult plants of O. grandiglumis accessions showed enhanced internodal elongation with rising water level and their growth response closely resembled that of deepwater varieties of O. sativa with high floating capacity. On the other hand, when subjected to complete submergence, seedlings of O. grandiglumis accessions displayed reduced shoot growth and resumed normal growth after desubmergence, similar to the response of submergence-tolerant varieties of O. sativa. Neither SUB1A nor the SK genes were detected in the O. grandiglumis accessions. These results indicate that the O. grandiglumis accessions are capable of adapting successfully to flooding by activating two contrasting mechanisms as the situation demands and that each mechanism of adaptation to flooding is not mediated by SUB1A or the SK genes.     

The invasive plant Alternanthera philoxeroides benefits from clonal integration in response to defoliation

Many notorious alien invasive plants have the capacity for vigorous clonal growth, and clonal integration may contribute to their invasiveness in response to various disturbances. Here, it is hypothesized that clonal integration affects the growth, biomass allocation, physiology, and compensatory response of the alien invasive clonal plant Alternanthera philoxeroides when faced with defoliation. To test these hypotheses, a growth experiment was conducted to investigate the effect of clonal integration on the responses of A. philoxeroides to different levels of defoliation. Daughter ramets that had been grown with stolon connections that were either severed from or connected to the mother plant were subjected to four defoliation levels: 0 (control), 30% (mild), 60% (moderate) and 90% (heavy) removal of leaf tissue. Defoliation greatly decreased growth (total biomass, number of ramets and total stolon length) but increased the maximum quantum yield of photosystem II (F_v/F_m) of daughter ramets. Clonal integration significantly increased growth, F_v/F_m and contents of non-structural carbohydrates (soluble sugars and total non-structural carbohydrates) of A. philoxeroides, and these effects were larger under heavier defoliation. Moreover, clonal integration markedly reduced the shoot/root ratio of A. philoxeroides, and these effects tended to increase with increasing levels of defoliation. These results support our hypothesis that A. philoxeroides benefits from clonal integration in response to defoliation, suggesting that clonal integration may be closely related to the invasiveness of A. philoxeroides in natural habitats with frequent disturbances.     

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。     

Comparative gas exchange performance during the wet season of three Brazilian Styrax species under habitat conditions of cerrado vegetation types differing in soil water availability and crown density

We studied the influence of environmental conditions in different vegetation types of the Brazilian savanna (cerrado s.l.) on CO₂ assimilation and water use efficiency of Styrax ferrugineus, S. camporum and S. pohlii, which are morphologically similar but have distinct distribution patterns. We measured leaf gas exchange and water relations in adult and young plants. Adult plants of S. ferrugineus were measured in the cerrado s. str.; plants of S. camporum at the edge of cerradão, a woodland vegetation; and in a riparian forest those of S. pohlii. Eight-month-old young plants were planted in the cerrado s. str., at the edge and in the understory of the cerradão, and in the understory of the riparian forest. For young plants, the high light availability in the cerrado s. str. resulted in a threefold greater CO₂ assimilation rate (A) compared to the other sites. A of adult plants under full irradiation (1800 μmol photons m⁻² s⁻¹) was significantly lower in S. pohlii compared to S. ferrugineus and S. camporum. Although sufficient soil water was available at every site at the end of the wet season, the extreme high water content in the riparian forest soil led to increased stomatal conductance (gs) and transpiration rate (E) for young plants of S. camporum and S. pohlii, but not of S. ferrugineus. However, the large gs in S. pohlii and S. camporum young plants did not result in higher A rates. The differences may indicate some specialization of S. ferrugineus to best growth in open savanna and of S. pohlii in riparian forest habitats. The high water use efficiency of S. camporum young plants in the understory of cerradão seems not yet sufficient to emphasize that this species benefits from this particular habitat. Because, while water supply during the wet season is quite sufficient, soil water deficits and frequent fires can occur during the dry season and might modify strongly the habitat conditions for the three Styrax species.     

Photosynthetic acclimation is important for post-submergence recovery of photosynthesis and growth in two riparian species

Background and Aims

Concomitant increases in O₂ and irradiance upon de-submergence can cause photoinhibition and photo-oxidative damage to the photosynthetic apparatus of plants. As energy and carbohydrate supply from photosynthesis is needed for growth, it was hypothesized that post-submergence growth recovery may require efficient photosynthetic acclimation to increased O₂ and irradiance to minimize photo-oxidative damage. The hypothesis was tested in two flood-tolerant species: a C₃ herb, Alternanthera philoxeroides; and a C₄ grass, Hemarthria altissima. The impact of low O₂ and low light, typical conditions in turbid floodwater, on post-submergence recovery was assessed by different flooding treatments combined with shading.

Methods

Experiments were conducted during 30 d of flooding (waterlogging or submergence) with or without shading and subsequent recovery of 20 d under growth conditions. Changes in dry mass, number of branches/tillers, and length of the longest internodes and main stems were recorded to characterize growth responses. Photosynthetic parameters (photosystem II efficiency and non-photochemical quenching) were determined in mature leaves based on chlorophyll a fluorescence measurements.

Key Results

In both species growth and photosynthesis recovered after the end of the submergence treatment, with recovery of photosynthesis (starting shortly after de-submergence) preceding recovery of growth (pronounced on days 40–50). The effective quantum yield of photosystem II and non-photochemical quenching were diminished during submergence but rapidly increased upon de-submergence. Similar changes were found in all shaded plants, with or without flooding. Submerged plants did not suffer from photoinhibition throughout the recovery period although their growth recovery was retarded.

Conclusions

After sudden de-submergence the C₃ plant A. philoxeroides and the C₄ plant H. altissima were both able to maintain the functionality of the photosynthetic apparatus through rapid acclimation to changing O₂ and light conditions. The ability for photosynthetic acclimation may be essential for adaptation to wetland habitats in which water levels fluctuate.Key words: Aerenchyma, Alternanthera philoxeroides, flooding, growth, Hemarthria altissima, low light, photosynthesis, shade, submergence, waterlogging, wetland plant     

塔里木荒漠河岸林干扰状况与林隙特征

对塔里木河中游荒漠河岸林林隙基本特征和干扰状况进行了研究。结果表明：形成的林隙形状近似于椭圆形,椭圆长短轴比率在扩展林隙(EG)和冠空隙(CG)有所不同,平均分别为1.52和2.31;林隙密度约为62.5个?hm-2, EG和CG在塔里木荒漠河岸林景观中的面积比例分别为69.52%和29.03%,干扰频率分别为1.45%?a-1和0.61%?a-1,林隙干扰返回间隔期约为164a。林隙大小结构表现出以小林隙为主的偏正态分布,EG大小40—200m2,CG大小0—80m2。林隙形成速率为1.30个?hm-2a-1,20—30a前形成的林隙最多。林隙形成方式由树木折干 枯立形成的最为普遍,占形成木总数95.73%。林隙大多由2—5株形成木形成, 而由4株形成木创造的林隙最多,平均每个林隙拥有形成木4.1株。林隙形成木主要为森林建群种,林隙形成木分布最多的径级在5—25cm,高度在4—8m,每株形成木所能形成的EG面积为27.12m2, CG面积为11.32m2。边缘木的径级结构呈正态分布,而高度结构呈偏左的正态分布,平均每个林隙拥有8.375株边缘木,林隙边缘木平均胸径比形成木平均胸径高73.1%,表明荒漠河岸林林隙干扰十分频繁,地下水位的持续下降是林隙形成的驱动力。     

Contrasting physiological responses by cultivars of Oryza sativa and O. glaberrima to prolonged submergence

Background and Aims Oryza glaberrima

is widely grown in flood-prone areas of African river basins and is subject to prolonged periods of annual submergence. The effects of submergence on shoot elongation, shoot biomass, leaf area and CO₂ uptake were studied and compared with those of O. sativa.

Methods

A wide selection of lines of O. sativa and O. glaberrima, including some classified as submergence tolerant, were compared in field and pot experiments. Plants were submerged completely for 31 d in a field experiment, and partially or completely for 37 d in a pot experiment in a growth chamber.

Key Results

Leaf elongation and growth in shoot biomass during complete submergence in the field were significantly greater in O. glaberrima than in O. sativa. So-called submergence-tolerant cultivars of O. sativa were unable to survive prolonged complete submergence for 31–37 d. This indicates that the mechanism of suppressed leaf elongation that confers increased survival of short-term submergence is inadequate for surviving long periods underwater. The O. sativa deepwater cultivar ‘Nylon’ and the ‘Yélé1A’ cultivar of O. glaberrima succeeded in emerging above the floodwater. This resulted in greatly increased shoot length, shoot biomass and leaf area, in association with an increased net assimilation rate compared with the lowland-adapted O. sativa ‘Banjoulou’.

Conclusions

The superior tolerance of deepwater O. sativa and O. glaberrima genotypes to prolonged complete submergence appears to be due to their greater photosynthetic capacity developed by leaves newly emerged above the floodwater. Vigorous upward leaf elongation during prolonged submergence is therefore critical for ensuring shoot emergence from water, leaf area extension above the water surface and a subsequent strong increase in shoot biomass.Key words: Flooding, leaf area, net assimilation rate, Oryza glaberrima, O. sativa, photosynthesis, rice, stress adaptation, submergence escape     

The influence of abiotic stress and phenotypic plasticity on the distribution of invasive Alternanthera philoxeroides along a riparian zone

Relatively few studies have compared invasibility and species invasiveness among microhabitats within communities, synchronously. We surveyed the abundance and performance of non-native Alternanthera philoxeroides (Mart.) Griseb. (alligator weed), its co-occurring native congener, Alternanthera sessilis (L.) DC. (sessile joyweed), and other species in a wetland community along a riparian zone in southeast China to test the hypotheses that: i) degree of invasion differs between different types of microhabitats within the community; and ii) microhabitat types that differ in invasibility also differ in soil resource availability or in sediment characteristics likely to affect resource availability; iii) phenotypic plasticity of A. philoxeroides may play a key role in its adaptation to diverse habitats as can be concluded from its extremely low genetic diversity in China. The study riparian zone comprises different types of microhabitats including wet abandoned field, swamp, marsh dunes and gravel dunes. Consistent with these hypotheses, cover of A. philoxeroides was high in abandoned fields (73 ± 2.9%) and swamps (94 ± 1.3%), which had high soil nutrients and water availability. On the contrary, cover of native A. sessilis was relatively high in marsh dunes and grave dunes, which had coarse gravel surfaces, low soil nutrients and low water availability. A. philoxeroides showed greater morphological plasticity in response to habitat variation. In abiotically harsh habitats, stems had limited growth, and were prostrate with weak adventitious roots at nodes, forming thin, scattered patches. In the two richer habitats, the highly branched plants spread over the water or soil surface, supporting dense stronger leaf-bearing stems which grew vertically. The growth pattern of A. sessilis among microhabitats did not exhibit significant variations. These results suggest that morphological plasticity and microhabitat types with high soil resources may facilitate invasions of A. philoxeroides.     

Palm live aboveground biomass in the riparian zones of a forest in Central Amazonia

Aboveground biomass estimates in the Amazon region remain uncertain, partly due to extrapolations based mainly on samples collected in upland terrains of terra-firme forests. Most biomass estimates were focused on dicotyledonous trees or included other plant groups as a category of trees. Palms dominate areas that represent 20% of the Brazilian Amazon. However, their contribution to biomass estimates and the variation within riparian zones remain poorly documented. We estimated the biomass of palms larger than 1–cm diameter at breast height (1.3 m aboveground) in riparian plots (n = 40); investigated the potential bias caused by the use of dicotyledonous- or family- rather than species-level equations for biomass estimation; compared palm biomass between riparian and non-riparian plots (n = 72); and evaluated the effects of soil, topography, and stream characteristics in riparian plots on palm biomass. Mean palm biomass in riparian zones estimated with species-level equations (27.50 ± 12.94 Mg/ha, range: 3.32–63.27 Mg/ha) was three times greater than biomass estimated with a family-level equation (9.04 ± 4.29 Mg/ha, range: 1.51–21.25 Mg/ha) and was greater than mean biomass estimated with a pantropical equation (20.46 ± 9.29 Mg/ha, range: 3.67–47.99 Mg/ha). Mean palm biomass in riparian zones was four times greater than in non-riparian zones. Palm biomass was high in flatter areas with poorly drained soils, but lower around streams with higher discharge. Inclusion of palms can contribute to reducing the uncertainties in biomass estimates in Amazonian forests. Recognition of the importance of riparian zones may improve conservation policies. Abstract in Portuguese is available with online material.     

Effects of submergence on the growth of Phragmites australis seedlings

Colonisation by reed seedlings, Phragmites australis (Cav.) Trin. ex Steud. is rare and usually occurs after drawdown and when shallow water prevails. P. australis seeds have high rates of germination but successful colonisation is dependent upon subsequent water depths. We investigated the capacity of young reed plants to resist a 4 weeks submergence stress within a 5 months period, and their subsequent recovery. A pond experiment examined the interactions between submergence depth and the age of the seedlings at submergence. Four submergence treatments were used. In two partial submergence treatments, 50 and 80% of the initial leaf area was submerged. In two total submergence treatments, plants were either submerged at 125% of their initial height with possible subsequent development of emerged leaves, or the water was deepened as they grew to maintain total submergence for 4 weeks. The ages at submergence were 40, 60 and 80 days. Plants were harvested at 5 months. Shoot elongation, biomass allocations to aerial biomass, roots and rhizomes, and photosynthetic activity of aerial leaves were measured. Redox potential was measured for a subsample.Mortality (18.7%) occurred only in the permanent submergence treatment for 40-day-old seedlings. In all treatments, submerged leaves senesced, except the terminal (youngest) leaves of permanently submerged plants. Submergence differentially affected shoot length and biomass, depending upon the intensity of the treatment and the seedling age. The major differences were found between the two partial and two total submergence treatments. Partial submergence (50 and 80%) significantly enhanced biomass accumulation and growth, whereas total submergence largely decreased biomass production and growth in length, with less effect on shoot numbers. The 80-day-old seedlings tolerated submergence better but growth was poorest in medium-aged plants (60-day-old). Increased elongation of the growing internodes of up to 140% was caused by submergence, and photosynthetic activity was enhanced by 85% in emergent leaves of plants initially submerged but allowed to produce emerged leaves during the treatment period.Young P. australis plants require shallow water levels without long lasting submergence to grow and survive. Tolerance to submergence increases with age. These processes contribute to define the conditions for colonisation via seeds in P. australis.