Reproductive allocation of biomass and nitrogen in annual and perennial Lesquerella crops

* BACKGROUND AND AIMS: The use of perennial crops could contribute to increase agricultural sustainability. However, almost all of the major grain crops are herbaceous annuals and opportunities to replace them with more long-lived perennials have been poorly explored. This follows the presumption that the perennial life cycle is associated with a lower potential yield, due to a reduced allocation of biomass to grains. The hypothesis was tested that allocation to perpetuation organs in the perennial L. mendocina would not be directly related to a lower allocation to seeds. * METHODS: Two field experiments were carried on with the annual Lesquerella fendleri and the iteroparous perennial L. mendocina, two promising oil-seed crops for low-productivity environments, subjected to different water and nitrogen availability. * KEY RESULTS: Seed biomass allocation was similar for both species, and unresponsive to water and nitrogen availability. Greater root and vegetative shoot allocation in the perennial was counterbalanced by a lower allocation to other reproductive structures compared with the annual Lesquerella. Allometric relationships revealed that allocation differences between the annual and the perennial increased linearly with plant size. The general allocation patterns for nitrogen did not differ from those of biomass. However, nitrogen concentrations were higher in the vegetative shoot and root of L. mendocina than of L. fendleri but remained stable in seeds of both species. * CONCLUSIONS: It is concluded that vegetative organs are more hierarchically important sinks in L. mendocina than in the annual L. fendleri, but without disadvantages in seed hierarchy.     

铅污染土壤中杂草对铅的吸收

利用盆栽试验,研究了杂草对铅的吸收积累及铅污染土壤中杂草养分吸收和根系丛枝菌根真菌(AMF)侵染的特点．结果表明,铅污染对杂草生长没有明显影响．铅主要富集在根系中,对某些杂草吸收养分有影响．苗期除了升马唐钾含量铅处理与对照之间有显著差异外,其它差异均不显著,说明苗期铅处理对鸡眼草、苦荬菜、升马唐和无芒稗吸收养分基本没有影响;成熟期铅处理中苦荬菜和升马唐的全磷含量极显著高于对照,而无芒稗全氮含量显著低于对照．根系AMF的侵染特点是,无论是苗期还是成熟期。鸡眼草和升马唐根系的AMF侵染率变化不大;苦荬菜铅处理低于对照,苗期铅处理和对照的根系侵染率分别是45．52％和69．44％,成熟期分别是74．64％和82．21％;无芒稗铅处理根系的AMF侵染率显著高于对照,苗期铅处理和对照的根系侵染率分别是82．45％和59．19％,成熟期分别是91．36％和78．28％．     

Partitioning and mobilization of starch and N reserves in grapevine (Vitis vinifera L.)

We followed C and N reserves of grapevines grown in trenches under semi-controlled conditions over a 3-year period after planting. Temporal mobilization of stored C and N and subsequent distribution of reserve materials within the vines were described in parallel with 15N uptake, particularly during the third growing season. Storage C in the perennial tissues (roots, trunk, canes) was mainly made of starch, which accumulated in the ray parenchyma of the wood. In the permanent tissues, starch and total nitrogen contents were found to decrease early in the development (bleeding sap, budbreak) whereas, on a concentration basis, they decreased only after stage 7 (first leaf fully expanded). Starch started to accumulate again in the perennial tissues during flowering. The same observation was made with total nitrogen, although N levels were much lower than those of starch. The 15N study showed that N uptake by the roots started at budbreak and increased with vine development, becoming predominant over reserve mobilization only after the onset of flowering. Taken together, these results indicate that the spring growth period can be divided into three main phases: In the first (dormancy to budbreak), significant losses of C and N proceed mainly via root necrosis. In the second period (first leaf to the onset of bloom), a strong mobilization of starch (and, to a lower extent, of N) occurred for supporting vegetative and reproductive growth. At that point, most of the C and N reserves used on the spring flush were those of the roots, rather than those of the old wood (trunk, canes). In the third period (bloom and early berry development), the mobilization process became low and was relieved by N uptake (and CO2 assimilation) supplying nutrients to the sink structures.     

Alternate Bearing Affects Nitrogen, Phosphorus, Potassium and Starch Storage Pools in Mature Pistachio Trees

The relationship between crop load and the functional storageof selected macronutrients and starch was assessed to developnutrient budgets and best management fertilization practicesin orchards. Functional storage represents the amount of nutrientsand starch redistributed from perennial tree parts in supportof the spring growth flush. Functional storage was influencedby:(a)nutrient and starch accumulation prior to dormancy; and(b)nutrientand starch demand by vegetative and reproductive organs in spring.Lightly cropping (off-year) trees stored 7, 14 and 2 times asmuch N, P and K, respectively, as heavily cropping (on-year)trees. Similar to many biennial plant species, nutrients thataccumulated during the vegetative phase in off-year trees wereused to support reproductive growth during the subsequent on-year.Soil nutrient uptake contributed more to storage pools thanleaf nutrient resorption in off-year-trees, while the reversewas true in on-year trees. Net nutrient resorption from senescingleaves accounted for all of the N and P and a third of the Kstored in on-year trees. Only between 20–33% of the N,P and K stored in perennial tissues of off-year trees couldbe attributed to leaf nutrient resorption. This is the firststudy to determine the amounts of nutrients stored in the perennialparts of mature, field-grown trees and the relative contributionsof leaf nutrient resorption and soil nutrient uptake to functionalstorage in trees.Copyright 1998 Annals of Botany Company Pistacia vera, nutrient storage, biennial bearing, crop load, leaf nutrient resorption, source-sink relationships.     

新疆典型盐生植物营养器官盐分积累与生态化学计量特征

对盐生植物体内器官盐分积累和碳(C)、氮(N)、磷(P)化学计量特征的研究, 有助于了解盐生植物养分、盐分的分配机制和其对盐渍环境的适应策略。该文选择新疆4种典型的真盐生植物(3种灌木植物: 盐节木(Halocnemum strobilaceum)、小叶碱蓬(Suaeda microphylla)、盐爪爪(Kalidium foliatum), 1种草本植物: 盐角草(Salicornia europaea)为研究对象, 对比研究了它们的根、茎、叶中的盐分积累和C、N、P化学计量特征以及二者间的相关性。结果显示: 1)在生长旺盛期, 4种盐生植物体内盐分积累可形成“盐岛”效应(Na⁺、Cl^-和电导率随盐离子从植物根部向顶端运输过程呈显著增加的趋势)。2) 4种盐生植物中灌木群落的生长主要受到P的限制; 草本群落的生长受N和P (偏P)共同限制。3)器官、物种和二者的交互作用均能影响盐生植物的盐分(除Mg²⁺外)特征和C、N、P生态化学计量特征。4) 4种盐生植物C与N、P之间显著负相关, N与P之间显著正相关。5)盐生植物体内总盐分、Na⁺和K⁺与N、N:P之间呈显著正相关关系, 与C、C:N、C:P之间呈显著负相关关系, 而K⁺和CO₃^2-与P之间却显著正相关。盐生植物体内盐分和养分在应对盐渍环境上存在一定的相互促进关系, 增施N肥有助于盐生植物对氯化物盐渍土的脱盐。这些发现为了解盐渍生境中盐生植物的生理生态适应及盐渍土的改良提供一定的理论依据。     

Salt accumulation in vegetative organs and ecological stoichiometry characteristics in typical halophytes in Xinjiang,China

AimsStudying salt accumulation in vegetative organs and carbon (C), nitrogen (N), phosphorus (P) stoichiometry in halophytes contributes to understand the adaptive strategy to saline environment and the distribution mechanism of nutrients and salinity of the halophytes.MethodsFour typical dominant species, including three perennial shrubs (Halocnemum strobilaceum, Suaeda microphylla and Kalidium foliatum) and an annual herb (Salicornia europaea) were selected for analyzing the salt accumulation and C, N, P stoichiometry relationships.Important findings 1) The salt accumulation showed “salt island” effect at peak growing stage; Na⁺, Cl^- and electrical conductivity all showed significant positive trends, when salt ions were transported from the root to the shoot of 4 halophytes. 2) P was the limiting growth element of perennial shrubs, while both N and P were limiting factors for the annual herb. 3) Different organs, species and their interactions affected C, N, P stoichiometry and salt icons except Mg²⁺. 4) C was negatively correlated with N and P, but there was a significant positive correlation between N and P. 5) Electrical conductivity, Na⁺and K⁺ were positively correlated with N, N:P ratio, and negatively correlated with C, C:N and C:P ratios, and yet K⁺, CO₃^2- had significant positive correlation with P. These results implied that there were some kind of mutual promoting relationships between nutrients and salinity in halophytes in the saline environment, and fertilizing with N could promote sodium chloride desalination.     

Distribution and levels of phytoecdysteroids in plants of the genus Silene during development

The purpose of the present work is the study of ecdysteroid distribution in annual and perennial Silene species during development. The experimental approach included the measurement of ecdysteroid levels in different plant organs and an evaluation of the contribution of individual organs to the total amount of 20-hydroxyecdysone (20E) produced by the plant. The highest concentrations of 20E were observed in reproductive organs. High levels were also found in leaves with lower levels in stems. Maximal ecdysteroid content for aerial parts was observed during periods of intense growth: at budding or flowering in annual species, and during vegetative growth or budding in perennial species. The contribution of the different organs to the overall ecdysteroid content changes during plant development. Leaves represent the main part of plant mass and 20E content. Reproductive organs represent a relatively small mass, but they contain high concentrations of ecdysteroids and, in terms of the amount of 20E they contain, their contribution is equal to that of stems. Arch.     

黄土高原退耕地的生态恢复

在刈割干扰下,黄土高原退耕地恢复进程中植被从沙蓬（Agriophyllum arenarium)单优群落向以达乌里胡枝子（Lespedeza davurica)为优势种的群落演替。物种增加速度初期快,后期慢,退耕后恢复初期的新增物种主要是沙蓬和白草（Pennisetum flaccidum);中前期主要是茵陈蒿（Artemisia capillaris),可成为群落的次优势种;中后期新增物种主要是多年生豆科植物,只是群落的伴生种,群落中1年生和多年生植物地上生物量分别以8.8%的年平均速率减少或增加,豆科植物地上生物量及所占比例上升,在退耕地恢复过程中,上层土壤中植物地下生物量比例与变化幅度增加,表现出浅层化趋势,退耕地恢复演替1-7年,土壤粘粒和粉粒减少,砂粒增加,7年后变化趋势相反,退耕地0-100cm土壤含水量在恢复期间呈逐渐上升趋势,中期增幅显著,全N和速效N在恢复前期减少,后期增加,在刈割利用下,退耕地恢复过程中土壤全P,速效P和有机C持续衰竭,退耕地恢复到当前稳定的群落所需时间分别为优势种群8-9年,群落9-11年和土壤11-12年。     

Species attributes in early primary succession on volcanoes

Abstract. We predicted that plants that can establish on volcanic soils with similar disturbance histories will have similar growth characteristics. We tested this prediction by a multivariate analysis of 27 traits of 84 species found six years after an eruption on Mount St. Helens, Washington State, USA, and Mount Usu, Hokkaido, Japan. These traits include vegetative, life-history, phenological and seed-biology characteristics. Cluster analysis revealed five species groups: annual herbs, perennial forbs, graminoids, shrubs and trees. Each group has distinct vegetative, life-history, and seed-biology traits. Except for shrubs, which were lacking on Mount Usu, both floras were well represented in each group. On intensely disturbed sites on both volcanoes, perennial forbs, whose development is dependent primarily on well-developed below-ground organs and wind-dispersal, expanded their cover more rapidly than did graminoids. These graminoids generally produce gravity-dispersed seeds and have close-set rhizomes and/or shoots. These results suggest that species that can establish during the early stages of succession on each volcano have similar vegetative, life-history, and seed-biology traits.     

新疆郁金香营养生长、个体大小和开花次序对繁殖分配的影响

植物有性繁殖与资源分配的关系研究对于揭示植物生活史特征及繁育系统进化具有重要意义。新疆郁金香(Tulipa sinkiangensis)是新疆天山北坡荒漠带特有的一种多年生早春短命植物。在自然生境中,该物种仅以有性繁殖产生后代,每株能产生1-8朵花,且不同植株上的花数及果实数以及花序不同位置上的花与果实大小明显不同。本文通过对新疆郁金香有性繁殖与营养生长及植株大小的关系以及花序中不同位置花及果实间的资源分配研究,旨在揭示营养生长、个体大小及开花次序对其繁殖分配的影响。结果表明:在开花和果实成熟阶段,新疆郁金香植株分配给营养器官(鳞茎和地上营养器官)与繁殖器官的资源间均存在极显著的负相关关系(P<0.01),说明其植株的营养生长与生殖生长间存在权衡关系。多花是新疆郁金香的一个稳定性状,其植株上花数目、花生物量、果实生物量和种子数量与植株生物量间均呈极显著的正相关关系(P<0.01),说明新疆郁金香植株的繁殖分配存在大小依赖性。在具2-5朵花的新疆郁金香植株中,花序内各花的生物量、花粉数和胚珠数、结实率、果实生物量、结籽数、结籽率及种子百粒重按其开花顺序依次递减,说明花序内各花和果实的资源分配符合资源竞争假说。植株通过减少晚发育的花或果实获得的资源来保障早发育的花或果实获得较多的资源,从而达到繁殖成功。     

Genetics of flower initiation and development in annual and perennial plants

Flowering is an integral developmental process in angiosperms, crucial to reproductive success and continuity of the species through time. Some angiosperms complete their life cycle within a year (annual plants), and others have a longer reproductive life, which is characterized by the generation of new flowering and vegetative shoots every year (perennial plants). Despite the differences in their lifespan, the underlying genetics of flower induction and floral organ formation appears to be similar among these plants. Hence, the knowledge gained from the study of flowering mechanism in Arabidopsis thaliana can be used to better understand similar processes in other plant species, especially the perennials, which usually have a long generation time and are not amenable to genetic analysis. Using Arabidopsis as a model, we briefly discuss the current understanding of the transition from vegetative to reproductive growth and the subsequent formation of individual floral organs, and how this knowledge has been successfully applied to the identification of homologous genes from perennial crops. Although annuals appear to share many similarities with perennials in terms of gene function, they differ in their commitment to flowering. Once an annual reaches the reproductive phase, all meristems are typically converted into either floral or inflorescence meristems. In contrast, each year, each meristem of a mature perennial has the choice to produce either a vegetative or a reproductive shoot. The physiology and genetics of flowering in Citrus are used to highlight the complexity of reproductive development in perennials, and to discus possible future research directions.     

Biomass and nutrient changes in the herbaceous layer of two central Australian mulga shrublands after unusually high rainfall

The amount of standing plant material in the ground storey and its botanical composition are compared for two central Australian mulga communities, one with a predominantly annual understorey and the other with a predominantly perennial understorey. Amounts of litter and nutrient levels in standing plant material, litter and soils are also compared. The number of species, particularly forbs, was greater in the annual community and differences in botanical composition are unlikely to have been caused by cattle grazing. The differences may be attributable in part to the greater proximity of the annual community to sources of alluvial material, and thus to better supplies of nutrients. For the 13 1/2 month study period following exceptional summer rain, the weight of herbage in the perennial community generally exceeded that in the annual community, whereas the weight of litter was less in the perennial community. Of a large number of plant and soil nutrients, phosphorus, calcium and magnesium were lower in the perennial community; these three nutrients, together with nitrogen, may be deficient for either plant growth or cattle diet in that community. The amount of sulphur was lower in the annual community but was unlikely to be deficient.     

Mineral allocation to reproduction in Sorhum bicolor and Sorghum halepense in relation to parental nutrient supply

Summary This experiment investigated the effect of parental nutrient shortage on the allocation of five nutrients to seeds and rhizomes in Sorghum halepense, a perennial, noxious weed, and to seeds in Sorghum bicolor, an annual, cultivated species. Plants from both species were grown from seeds and supplied with fertilizer at three concentrations. The allocation of biomass and nutrients (N, P, K, Ca and Mg) to reproductive and vegetative parts was determined. Relative biomass allocation to reproduction (either sexual or vegetative) remained constant in S. halepense in spite of large differences in total plant weight. In S. bicolor, however, biomass allocation to sexual reproductive structures decreased significantly with decreasing nutrient supply. Individual seed weight was not modified by parental nutrient supply in S. halepense, but it increased with decreasing nutrient availability in S. bicolor. Important differences in mineral allocation to seeds were found between the two species. While S. bicolor seeds were largely buffered from the differences in parental nutrient status, concentration of nutrients in S. halepense seeds decreased significantly with decreasing supply for all the nutrients analyzed except Ca. However, mineral nutrient concentration in S. halepense rhizomes remained remarkably constant despite differences in the external supply, evincing the priority given to vegetative reproduction at the expense of sexual reproduction. Overall, the pattern of nutrient allocation in S. bicolor seeds under different nutrient supply resembled the pattern observed in S. halepense rhizomes, but it had little resemblance to the pattern of nutrient allocation in S. halepense seeds. The results are discussed in terms of differences and similarities in the reproductive strategy of these two species.     

Adaptive genetic differentiation in life-history traits between populations of <Emphasis Type="Italic">Mimulus guttatus</Emphasis> with annual and perennial life-cycles

The optimal allocation to sexual and vegetative reproduction as well as the optimal values of other life-history characteristics such as phenology, growth and mating system are likely to depend on the life-cycle of the organism. I tested whether plants of Mimulus guttatus originating from temporarily wet populations where the species has an enforced annual life-cycle have higher allocation to sexual reproduction, lower allocation to vegetative reproduction, more rapid phenology, faster growth, and floral traits associated with a self-fertilizing mating system than plants from permanently wet populations where the species has a perennial life-cycle. I grew a total of 1377 plants originating from three populations with an annual life-cycle and 11 populations with a perennial life-cycle in a greenhouse under permanently and temporarily wet conditions. Plants of M. guttatus in permanently wet conditions had significantly more vegetative reproduction and tended to have a faster growth than plants in the temporarily wet conditions, indicating plasticity in these life-history traits. Plants from populations with an annual life-cycle invested significantly more in sexual reproduction and significantly less in vegetative reproduction than the ones from populations with a perennial life-cycle. Moreover, this study showed that plants originating from populations with an annual life-cycle have a significantly faster development and floral traits associated with autonomous self-fertilization. In conclusion, this study suggests that there has been adaptive evolution of life history traits of M. guttatus in response to natural watering conditions that determine the life span of the species.     

miRNA control of vegetative phase change in trees

After germination, plants enter juvenile vegetative phase and then transition to an adult vegetative phase before producing reproductive structures. The character and timing of the juvenile-to-adult transition vary widely between species. In annual plants, this transition occurs soon after germination and usually involves relatively minor morphological changes, whereas in trees and other perennial woody plants it occurs after months or years and can involve major changes in shoot architecture. Whether this transition is controlled by the same mechanism in annual and perennial plants is unknown. In the annual forb Arabidopsis thaliana and in maize (Zea mays), vegetative phase change is controlled by the sequential activity of microRNAs miR156 and miR172. miR156 is highly abundant in seedlings and decreases during the juvenile-to-adult transition, while miR172 has an opposite expression pattern. We observed similar changes in the expression of these genes in woody species with highly differentiated, well-characterized juvenile and adult phases (Acacia confusa, Acacia colei, Eucalyptus globulus, Hedera helix, Quercus acutissima), as well as in the tree Populus x canadensis, where vegetative phase change is marked by relatively minor changes in leaf morphology and internode length. Overexpression of miR156 in transgenic P. x canadensis reduced the expression of miR156-targeted SPL genes and miR172, and it drastically prolonged the juvenile phase. Our results indicate that miR156 is an evolutionarily conserved regulator of vegetative phase change in both annual herbaceous plants and perennial trees.     

Environmental heterogeneity and mechanism of stoichiometry properties of vegetative organs in dominant shrub communities across the Loess Plateau

 为了探究黄土高原灌丛群落中优势物种的根、茎和叶等营养器官之间碳(C)、氮(N)、磷(P)及其比值等化学计量特征的环境分异性及其与土壤养分的耦合性, 在甘肃省和宁夏回族自治区境内的3个灌丛集中分布区(甘肃南部、宁夏北部和甘肃西部)沿水热梯度选取41个样点进行样地调查。结果显示: 1)甘肃、宁夏灌丛群落的有机物质含量及P资源相对匮乏, 而N资源相对丰富。2)从甘肃南部、宁夏北部到甘肃西部, 生长季温度递增、年降水量递减, 与此耦合, 土壤养分也逐级递减, 沿着土壤养分梯度, 黄土高原优势灌丛根、茎和叶的C、N、P储量减少, 根和茎的C:N下降, 根、茎和叶的N:P上升, 但在宁夏北部和甘肃西部间差异不显著。同时, 3个优势灌丛分布区的优势灌丛各器官间营养元素的分配格局不同。3)土壤养分相对较高的区域优势灌丛间各器官营养元素储量无差异, 而土壤养分较低区域亲缘关系较远的优势灌丛间各器官的营养元素储量差异显著, 而亲缘关系较近的优势灌丛各器官营养元素储量差异不显著。黄土高原优势灌丛各器官C、N、P化学计量特征是植物体与土壤中化学元素耦合的结果, 当土壤养分逐渐升高时, 植物体内的化学元素储量也逐渐增多。该研究不仅有助于认识黄土高原优势灌丛化学计量环境分异规律, 而且有助于洞察不同土壤条件下C、N、P在优势灌丛营养器官间的分配格局和植物资源分配策略, 并为黄土高原植被的管理和恢复提供一定的理论基础。     

Variation in resource abundance affects diet and feeding morphology in the pumpkinseed sunfish (Lepomis gibbosus)

Summary Growth responses and accumulation of N and P were studied in two pygmy south-west Australian species of Drosera following supplementary feeding of arthropods (collembolans, Hypogastrura vernalis and fruit flies, Drosophila melanogaster) and/or a balanced mineral nutrient supplement (N as nitrate) via the roots. One feeding experiment used glasshouse-raised germlings from vegetative propagules (gemmae) of the perennial Drosera closterostigma, the other three (two on D. closterostigma and one on the annual D. glanduligera) involved natural populations engaging in natural captures of indigenous prey. All experiments recorded highly significant increases in plant dry matter, N and P (all plant age groups) and in reproductive performance (adult plants only) from artificial feeding of arthropods, but no apparent benefits from minerals alone or additive effects of minerals above that due to insects. Unresponsiveness to mineral nutrients was suggested to relate to inability of the species to use nitrate, while up to three-fold growth and nutrient uptake response to insects indicated that growth of natural populations might be severely limited by inadequate catches of prey. It is concluded that the highly nutrient-poor conditions typical of the habitat of pygmy species of Drosera may have promoted marked specialization towards carnivory and an attendant decline in ability to utilize soil-derived sources of nutrients.     

Total uptake of nutrients at different stages of the growth of groundnut and the ratios in which various nutrient elements exist in groundnut plant

Summary A pot experiment was conducted with groundnut crop in two red soils with various levels of application of calcium, phosphorus, potassium and magnesium. The plants were analysed at vegetative, reproductive and post harvest stages for the uptake of nutrients. Up to vegetative stage, the crop absorbed 10 per cent of the total requirement of nitrogen, phosphorus, calcium and magnesium. During the reproductive stage the crop removed about 40 to 50 per cent of the above nutrients. Between reproductive and harvest stages groundnut absorbed the remaining amount of the nutrients.Forms a part of the Ph.D. thesis of the first author approved by Tamil Nadu Agricultural University.