Seasonal control over allocation to reproduction in a tussock-forming and a rhizomatous species of Eriophorum in central Alaska

Summary The evergreen tussock-forming Eriophorum vaginatum revealed consistently earlier (c. 1 moth) phenology and greater biomass per tiller than the summergreen rhizomatous E. scheuchzeri in all four components measured (vegetative and reproductive shoots and stems) under the same climatic regime in central Alaska over one growing season. Greatest allocation to vegetative shoot growth occurred in mid-summer in both species. The tussock growth form of E. vaginatum raised shoot meristems 25–30 cm above the soil surface, where temperatures were warmer, permitting shoot growth to begin earlier in spring and continue longer in autumn than in E. scheuchzeri. Consequently, E. vaginatum was able to allocate reserves to reproductive tillers primarily in autumn and early spring, times when minimal reserves were required for vegetative growth. By contrast, the rhizomatous E. scheuchzeri had a more constrained growing season, and allocation to reproduction coincided with allocation to vegetative growth. For this reason, reserves were drawn down more fully in mid-summer in E. scheuchzeri than in E. vaginatum. The more conservative use of nutrient stores in E. vaginatum may relate to its great longevity, reduced allocation to reproduction (including low seedling recruitment), and relatively stable habitats. The mid-seasonal pulse of allocation to reproduction in E. scheuchzeri appears viable only in relatively fertile disturbed sites, where the soil nutrient supply is sufficient to support simultaneous allocation to vegetative growth and reproduction.     

Biomass partitioning,architecture and turnover of six herbaceous species from habitats with different nutrient supply

Three grasses (Holcus lanatus, Anthoxanthum odoratum and Festuca ovina) and three herbs (Rumex obtusifolius, Plantago lanceolata and Hieracium pilosella) were grown in a greenhouse at 3 nutrient levels in order to evaluate plant allocation, architecture and biomass turnover in relation to fertility level of their habitats.Four harvests were done at intervals of 4 weeks. Various plant traits related to biomass partitioning, plant architecture, biomass turnover and performance were determined. Differences in nutrient supply induced a strong functional response in the species shoot:root allocation, but architecture and turnover showed little or no response. Architectural parameters like specific leaf area and specific root length, however, in general decreased during plant development.Species from more nutrient-rich successional stages were characterized by a larger specific leaf area and longer specific shoot height (height/shoot biomass), resulting in a higher RGR and total biomass in all nutrient conditions. There was no evidence that species from nutrient-poor environments had a longer specific root length or any other superior growth characteristic. The only advantage displayed by these species was a lower leaf turnover when expressed as the fraction of dead leaves and a shorter specific shoot height (SSH) which might prevent herbivory and mowing losses.The dead leaf fraction, which is a good indicator for biomass and nutrient loss, appeared to be not only determined by the leaf longevity, but was also found to be directly related to the RGR of the species. This new fact might explain the slow relative growth rates in species from a nutrient-poor habitat and should be considered in future discussions about turnover.     

Effects of light and nutrient availability on the growth,allocation, carbon/nitrogen balance,phenolic chemistry,and resistance to herbivory of two freshwater macrophytes

Phenotypic responses of Potamogeton amplifolius and Nuphar advena to different light (7% and 35% of surface irradiance) and nutrient environments were assessed with field manipulation experiments. Higher light and nutrient availability enhanced the growth of P. amplifolius by 154% and 255%, respectively. Additionally, biomass was allocated differently depending on the resource: high light availability resulted in a higher root/shoot ratio, whereas high nutrient availability resulted in a lower root/shoot ratio. Low light availability and high nutrient availability increased the nitrogen content of leaf tissue by 53% and 40% respectively, resulting in a 37% and 31% decrease in the C/N ratio. Root nitrogen content was also increased by low light and high nutrient availability, by 50% (P=0.0807) and 77% respectively, resulting in a 20% and 40% decrease in root C/N ratio. Leaf phenolics were significantly increased 72% by high light and 31% by high nutrient availability, but root phenolic concentrations were not altered significantly. None of these changes in tissue constituents resulted in altered palatability to crayfish. N. advena was killed by the same high nutrient treatment that stimulated growth in P. amplifolius, preventing assessment of phenotypic responses to nutrient availability. However, high light availability increased overall growth by 24%, but this was mainly due to increased growth of the rhizome (increased 100%), resulting in a higher root/shoot ratio. High light tended to increase the production of floating leaves (P=0.09) and significantly decreased the production of submersed leaves. High light availability decreased the nitrogen content by 15% and 25% and increased the phenolic concentration by 88% and 255% in floating and submersed leaves, respectively. These differences in leaf traits did not result in detectable differences in damage by herbivores.     

Über den Verzweigungsvorgang beiPsilotum undSelaginella mit Anmerkungen zum Begriff der Dichotomie

After a critical evaluation of the concept of dichotomous branching in Cormophytes the shoot apical meristems ofPsilotum triquetrum andSelaginella speciosa are described. InPsilotum only the terminal meristems of the cryptophilic shoots have a three sided apical cell. Those of the photophilic shoots lack a typical apical cell.Selaginella has a two sided apical cell. The process of branching is independent from apical cells. It is due to an equal or unequal fractionation of the initial zone of the shoot apex which embraces all tissues above the leaf producing zone of the apical meristem.

Herrn Univ.-Prof. Dr.Walter Leinfellner zum 70. Geburtstag gewidmet.     

Intra-specific variation in relative growth rate: impact on competitive ability and performance of Lychnis flos-cuculi in habitats differing in soil fertility

Plant species from unproductive or adverse habitats are often characterized by a low potential relative growth rate (RGR). Although it is generally assumed that this is the result of selection for specific trait combinations that are associated with a low rate of net biomass accumulation, few studies have directly investigated the selective (dis-)advantage of specific growth parameters under a set of different environmental conditions. Aim of the present study was to quantify the impact of inherent differences in growth parameters among phenotypes of a single plant species, Lychnis flos-cuculi, on their performance under different soil nutrient conditions. Growth analysis revealed significant variation in RGR among progeny families from a diallel cross between eight genotypes originating from a single population. Differences in RGR were due to variation in both leaf area ratio (LAR) and in net assimilation rate (NAR). A genetic trade-off was observed between these two components of growth, i.e. progeny families with high investment in leaf area had a lower rate of net biomass accumulation per unit leaf area. The degree of plasticity in RGR to nutrient conditions did not differ among progeny families. Inherent differences in growth parameters among progeny families had a significant impact on their yield in competition with Anthoxanthum odoratum and Taraxacum hollandicum. In nutrient-rich conditions, progeny families with an inherently high leaf weight ratio (LWR) achieved higher yield in competition, but variation in this trait could not explain differences in competitive yield under nutrient-poor conditions. Inherent differences in growth parameters among progeny families were poorly correlated with differences in survival and average rosette biomass (a good predictor of fecundity) among these progeny families sown in four field sites along a natural gradient of soil fertility. In the more productive sites none of the growth parameters was significantly correlated with rosette biomass, but in the least productive site progeny families with an inherently high specific leaf area (SLA) tended to produce smaller rosettes than low-SLA families. These results are consistent with the view that a selective advantage may accrue from either high or low values of individual RGR components, depending on habitat conditions, and that the selective advantage of low trait values in nutrient-poor environments may results in indirect selection for low RGR in these habitats.     

麻栎和闽楠幼苗叶功能性状及生物量对光照和施肥的响应

光照和养分条件是影响植物生长的重要环境因子,不同生活型植物对环境异质性的响应机制不同。以落叶阔叶树种麻栎和常绿阔叶树种闽楠幼苗为研究对象,设置2个光照梯度（全光照和45%全光照）和4个施肥梯度（不施肥、氮磷供应比为5、15和45）共8种处理,研究光照和施肥及其交互作用对麻栎和闽楠生物量和叶形态、生理及化学性状的影响,并探讨了叶功能性状和生物量的关系。结果表明：（1）光照、施肥及其交互作用对光合气体交换参数（除水分利用效率外）、叶绿素荧光参数、叶形态指标（除比叶面积外）、单位质量叶氮含量和根冠比影响显著（P<0.05）。此外,光照和施肥对地上生物量和总生物量影响显著（P<0.05）。（2）全光照显著增加了麻栎和闽楠单株总叶面积和地上、地下生物量及总生物量（P<0.05）,而遮荫降低了非光化学猝灭系数、光合氮利用率和根冠比,增加了单位质量叶氮含量。（3）在全光照处理中,施肥显著增加了麻栎和闽楠水分利用效率（P<0.05）;在遮荫处理中,氮磷供应比45显著增加了麻栎和闽楠净光合速率和水分利用效率（P<0.05）。（4）麻栎和闽楠在全光照中倾向于资源获取策略,在遮荫中偏向于资源保守策略。在光照和施肥处理中,麻栎和闽楠单株总叶面积与地上生物量均显著正相关（P<0.05）。总之,单株总叶面积是预测麻栎和闽楠幼苗地上生物量变化的稳定指标,施肥有助于增加低光环境下麻栎和闽楠幼苗的生态适应能力。     

<Emphasis Type="Italic">WUS</Emphasis> and <Emphasis Type="Italic">STM</Emphasis>-based reporter genes for studying meristem development in poplar

We describe the development of a reporter system for monitoring meristem initiation in poplar using promoters of poplar homologs to the meristem-active regulatory genes WUSCHEL (WUS) and SHOOTMERISTEMLESS (STM). When ~3 kb of the 5′ flanking regions of close homologs were used to drive expression of the GUSPlus gene, 50–60% of the transgenic events showed expression in apical and axillary meristems. However, expression was also common in other organs, including in leaf veins (40 and 46% of WUS and STM transgenic events, respectively) and hydathodes (56% of WUS transgenic events). Histochemical GUS staining of explants during callogenesis and shoot regeneration using in vitro stems as explants showed that expression was detectable prior to visible shoot development, starting 3–15 days after explants were placed onto callus inducing medium. A minority of WUS and STM events also showed expression in the cambium, phloem, or xylem of regenerated, greenhouse grown plants undergoing secondary growth. Based on microarray gene expression data, a paralog of poplar WUS was detectably up-regulated during shoot initiation, but the other paralog was not. Both paralogs of poplar STM were down-regulated threefold to sixfold during early callus initiation. We identified 15–35 copies of cytokinin response regulator binding motifs (ARR1AT) and one copy of the auxin response element (AuxRE) in both promoters. Several of the events recovered may be useful for studying the process of primary and secondary meristem development, including treatments intended to stimulate meristem development to promote clonal propagation and genetic transformation. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The effect of varying nitrogen and phosphorus availability on nutrient use by Larrea tridentata,a desert evergreen shrub

Summary In a phytotron study of the effects of nitrogen and phosphorus supply ratio on nutrient uptake and use by Larrea tridentata, seedlings responded to increases in N and P availability with increases in leaf size, total biomass, and leaf nutrient concentration, and with decreases in root: shoot ratio. N and P use efficiency decreased with increasing N and P availability, respectively, but increased with increasing availability of the other nutrient, suggesting that Larrea responds both to the absolute and to the relative availability of limiting nutrients. Absolute amounts of N and P resorption, as well as N and P resorption efficiencies did not demonstrate a significant trend with nutrient availability, and there was no evidence of significant interactions between the two nutrients. More studies of the effects of nutrient interactions in the cycling and use of nutrients by different plant species are needed before more general conclusions can be drawn.     

Morphological responses to nutrient availability in four clonal herbs

This paper examines morphological plasticity of clonal plants of contrasting habitats and of contrasting architectures in response to nutrient supply. The hypotheses were tested that plants from rich habitats possess greater plasticity in response to variation in resource supply than species from poor habitats, and that rhizomatous species are less plastic in their response than stoloniferous species. Two sympodial rhizomatous herbs (Carex flacca, C. hirta) and two monopodial stoloniferous herbs (Trifolium fragiferum, T. repens) were subjected to four levels of nutrient supply in a garden experiment. One of the two species of each genus (C. hirta, T. repens) is from fertile and the other from infertile habitats. We measured 1) whole plant characters: total plant dry weight, number of modules (product of a single apical meristem) and number of ramets; 2) ramet characters: ramet leaf area and ramet height; and 3) spacer characters: branches per module, length per module and length per module internode.All measured characters in the Trifolium species significantly responded to treatment: the values for all measured characters increased with higher levels of fertilization. The differences in plant characters between fertilization levels were larger in Trifolium repens than in T. fragiferum in terms of whole plant characters, ramet characters and stolon internode length. The two Carex species did not differ in their responses to treatment in terms of most characters measured. In ramet characters and in some whole plant characters the species from fertile habitats were more plastic than those from infertile habitats. In spacer characters this pattern was not found. Foraging could not be demonstrated unequivocally.Morphological plasticity in the stoloniferous (Trifolium) species was much larger than in the rhizomatous (Carex) species. This seems in accordance with a foremost storage function of rhizomes, as against a foremost explorative function of stolons.     

Allocation,within and between organs,and the dynamics of root length changes in two birch species

Spatial and temporal dynamics of biomass allocation within and between organs were investigated in seedlings of two birch species of contrasting successional status. Seedlings of Betula alleghaniensis Britt (yellow birch) and B. populifolia Marsh (gray birch) were grown for 6 weeks at two nutrient levels in rectangular plexiglass containers to allow non-destructive estimates of root growth, production and loss. Leaf area and production were simultaneously monitored. Yellow birch responded more to nutrient level than gray birch in terms of total biomass, shoot biomass, leaf area and root length. Yellow birch also flexibly altered within-organ allocation (specific leaf area, specific root length and specific soil amount). In contrast, gray birch altered between-organ allocation patterns (root length:leaf area and soil amount:leaf area ratios) more than yellow birch in response to nutrient level. Yellow birch showed greater overall root density changes within a very compact root system, while gray birch showed localized root density changes as concentric bands of new root production spread through the soil. Species differ critically in their responses of standing root length and root production and loss rates to nutrient supply. Early successional species such as gray birch are hypothesized to exhibit higher plasticity in varied environments than later successional species such as yellow birch. Our results suggest that different patterns of allocation, within and between plant organs, do not necessarily follow the same trajectories. To characterize thoroughly the nature of functional flexibility through ontogeny, within- and between-organ patterns of allocation must be accounted for.     

Development of maize plants from cultured shoot apices

Excised shoot apices of maize (Zea mays L.), comprising the apical meristem and one or two leaf primordia, have been cultured and can form rooted plantlets. The plantlets, derived from meristems that had previously formed 7–10 nodes, develop into mature, morphologically normal plants with as many nodes as seed-grown plants. These culture-derived plants exhibited the normal pattern of development, with regard to the progression of leaf lengths along the plant and position of axillary buds and aar shoots. Isolation of the meristem from previously formed nodes reinitiates the pattern and number of nodes formed in the new plant. Thus, cells of the meristem of a maize plant at the seedling stage are not determined to form a limited number of nodes.     

Shoot development and production studies of phragmites Australis (Cav.) Trin. Ex Steudel in Scottish Lochs

Studies were made on the development and production of Phragmites australis (Cav.) Trin. ex Steudel (= Phragmites communis Trin.) stands from Forfar Loch-polytrophic, Balgavies Locheutrophic and Loch of the Lowes-mesotrophic. Differences were detected in the shoot height, shoot dry weight, standing crop biomass, leaf area index, crop growth rate and net efficiency of solar energy conversion among the three lochs. Such differences were partly due to the different nutrient levels of the three locations. The percentage ratio of leaf to shoot weight, leaf area ratio, relative growth rate and net assimilation rate for the reed stands in the three lochs were also calculated and were compared with values from other study areas.     

不同生境下假臭草生长特征分析

通过测定庄稼地、弃耕地、桉树林、公路边4种生境下假臭草叶面积(LA)、比叶面积(SLA)、LDMC和株高等形态特征、植株各构件生物量及生物量分配比和假臭草种群盖度、密度等种群数量性状,研究不同生境类型下假臭草个体的形态特征、生物量及生物量分配和种群生长状况的差异。结果表明:光照和土壤养分对假臭草的形态特征、生物量及生物量分配和种群数量性状的影响显著;低光照、高土壤养分生境下假臭草的LA、SLA、株高较高,假臭草叶片的LDMC较小,植株的花序数较少,生物量主要向叶和茎器官进行分配,根和花果器官的生物量分配则受到限制;高光照、养分贫瘠生境下假臭草的LA、SLA、株高较低,假臭草叶片的LDMC较大,生物量主要向根和花果器官进行分配,种群的盖度、密度、生物量和高度等种群数量性状较低,种群生长状况较差;高光照、肥沃养分生境为假臭草的最适生境,假臭草构件的资源配置较为优化,株高、基径粗、冠宽和花序数等形态指标较大,种群盖度、密度、生物量等种群数量性状较大,种群生长较好。     

Production and nitrogen responses of the African dwarf shrub Indigofera spinosa to defoliation and water limitation

Summary The dwarf shrub Indigofera spinosa Forsk. (Papilionacea), a native forage species of arid Northwest Kenya, was propogated from seed, grown in a controlled environment, and subjected to three treatments of defoliation and watering frequencies in a factorial experimental design. Biomass production and nitrogen accumulation in tissue components were measured to determine defoliation responses in a water-limited environment. We hypothesized that plants would maintain biomass and nitrogen flows despite removal of aboveground meristems and tissues by defoliation. Principal experimental results included a slight reduction (11%; P=0.08) of total biomass production by clipping ca. 1/3 or 2/3 of new leaves and stems and all apical meristems every month. Total aboveground production was not affected by clipping, while final root biomass was reduced 17% by the 2/3 clipping. The least water stressed plants were affected most negatively by defoliation, and the unclipped plants responded more negatively to greater water limitation. Plants achieved partial biomass compensation through alterations in shoot activity and continued allocation of photosynthate to roots. A smaller fraction of leaf production was directed to litter in clipped plants although clipping only removed the youngest tissues, suggesting that clipping increased leaf longevity. In turn, each leaf probably contributed a greater total quantity of photosynthate. Photosynthetic rates were also likely to have been increased by clipping water-stressed plants. In contrast to biomass, plants overcompensated for nitrogen lost to defoliation. Total nitrogen uptake by individual plants was stimulated by defoliation, as there was more total nitrogen in leaves and stems. Increased nitrogen uptake was achieved by clipping stimulation of total uptake per unit of root rather than of total root mass.     

Transpiration by an emergent macrophyte: source of water and implications for nutrient supply

The contribution of sediment interstitial water and the water column to the transpiration stream of Myriophyllum aquaticum (Vellozo) Verdcourt was determined to estimate the significance of mass flow in supply of sediment nutrients for plant growth. Sediment interstitial water accounted for about 2% of the water transpired over a 37 day period. Because of the small volume of water that originated in the sediment we concluded that mass flow did not significantly enhance nutrient supply to the roots of M. aquaticum. Relative growth rate (RGR) of adventitious, water roots was greater than whole plant RGR, and RGR of sediment roots was not significantly different from zero, indicating a shift in the biomass allocation after emergence of the apical meristem into the air. Water use, measured by the transpiration coefficient, averaged 260 ml H₂O mg DW^-1, which is similar to C-4 terrestrial plants. M. aquaticum has leaf characteristics commonly associated with xerophytic habitats. These characteristics may be necessary if a high transpiration rate and a mechanical requirement for high cell turgor pressure, required by a reliance upon hydrostatic pressure for support of the aerial stems, are mutually exclusive because of morphological constraints on hydraulic conductivity.     

Adventitious shoot formation in decapitated dicotyledonous seedlings starts with regeneration of abnormal leaves from cells not located in a shoot apical meristem

Regeneration of new shoots in plant tissue culture is often associated with appearance of abnormally shaped leaves. We used the adventitious shoot regeneration response induced by decapitation (removal of all preformed shoot apical meristems, leaving a single cotyledon) of greenhouse-grown cotyledon-stage seedlings to test the hypothesis that such abnormal leaf formation is a normal regeneration progression following wounding and is not conditioned by tissue culture. To understand why shoot regeneration starts with defective organogenesis, the regeneration response was characterized by morphology and scanning electron and light microscopy in decapitated cotyledon-stage Cucurbita pepo seedlings. Several leaf primordia were observed to regenerate prior to differentiation of a de novo shoot apical meristem from dividing cells on the wound surface. Early regenerating primordia have a greatly distorted structure with dramatically altered dorsoventrality. Aberrant leaf morphogenesis in C. pepo gradually disappears as leaves eventually originate from a de novo adventitious shoot apical meristem, recovering normal phyllotaxis. Similarly, following comparable decapitation of seedlings from a number of families (Chenopodiaceae, Compositae, Convolvulaceae, Cucurbitaceae, Cruciferae, Fabaceae, Malvaceae, Papaveraceae, and Solanaceae) of several dicotyledonous clades (Ranunculales, Caryophyllales, Asterids, and Rosids), stems are regenerated bearing abnormal leaves; the normal leaf shape is gradually recovered. Some of the transient leaf developmental defects observed are similar to responses to mutations in leaf shape or shoot apical meristem function. Many species temporarily express this leaf development pathway, which is manifest in exceptional circumstances such as during recovery from excision of all preformed shoot meristems of a seedling.     

不同生境鱼腥草形态特征及主要有效成分含量差异分析

以不同生境中的鱼腥草为研究对象,分析其形态特征及主要有效成分含量的差异。该文测定了不同生境下主要环境因子指标及鱼腥草的叶长、叶宽、株高、基径、生物量和含水量,用高效液相色谱法测定其各部分主要有效成分金丝桃苷、槲皮苷含量,并对测量结果进行单因素方差分析和相关性分析。结果表明:环境因子会对鱼腥草的形态建成及有效成分积累造成不同影响,适宜的光照强度有利于鱼腥草对有效成分的累积;鱼腥草中有效成分的含量表现出叶茎根的规律。适宜的遮阴处理,既可避免强光照对鱼腥草生长的抑制,又可显著提高有效成分的含量,从而整体增加对鱼腥草有效成分的收益。