Heteroblasty and preformation in mayapple, Podophyllum Peltatum (Berberidaceae): developmental flexibility and morphological constraint

Developmental preformation can constrain growth responses of shoots to current conditions, but there is potential for flexibility in development preceding formation of the preformed organs. Mayapple (Podophyllum peltatum) is strongly heteroblastic, producing rhizome scales, bud scales, and either a single vegetative foliage leaf or two foliage leaves on a sexual shoot. To understand how and when preformation constrains growth responses, we compare (1) how leaf homologs of the renewal shoot differ in development, (2) whether there are differences in shoot development that occur in advance of morphological determination of shoot type, and (3) whether there are points of developmental flexibility in renewal shoot growth prior to preformation of the foliage and floral organs. We use scanning electron microscopy and histology to show that the three vegetative leaves (both types of scale leaves and the vegetative foliage leaf) are similar in the initial establishment of an encircling and overarching leaf base. Differences among them are found in the timing of differentiation of the leaf base and in the relative timing and degree of growth of the lamina and petiole. In contrast, foliage leaves on sexual shoots show less expression of the leaf base and precocious growth of the lamina and petiole. Prior to shoot type determination, there are no morphological differences in the sequence or position of leaf homologs that predict final shoot type. In this colony, leaves at positions 12 and 13, on average, appear to be identical in development until they are between 700 and 800 μm in length, when it becomes possible to distinguish leaves that will become vegetative foliage leaves from additional bud scale leaves on vegetative or sexual shoots. We suggest that late developmental determination of leaves at positions 12 and 13 reflects ontogenetic sensitivity to a transition to flowering. Thus, in mayapple, heteroblasty appears to facilitate developmental flexibility prior to the point where shoot growth becomes constrained by preformation of determined aerial structures.     

Morphology and development of floral shoots and organs in certain Zannichelliaeeae

Development of reproductive shoots and associated organs in Vleisia aschersoniana, Althenia filiformis, Lepilaena bilocularis and L. cylindrocarpa (Zannichelliaeeae, sensu Dumortier) has been examined using an epi-illumination technique to provide photographic documentation. Floral shoots share a similar basic developmental pattern. The vegetative shoot is terminated by a unisexual flower, but growth is continued from the axils of leaves immediately below so that fairly regular sympodia develop. Vleisia is most variable in the expression of this pattern. The flowers are simple, consisting of a single stamen or three carpels (one in Vleisia) and show marked similarity in development. Short scale-like appendages, reminiscent of a perianth, develop at the base of the stamen in Althenia and Lepilaena. An outgrowth at the tip of the connective in Lepilaena bilocularis and two at its middle part in Vleisia are initiated at late stages of stamen development. Carpels are subtended by membranous tepal-like appendages that are initiated at the same time as the carpel primordia. Each carpel primordium becomes peltate and develops a bitegmic ovule on the adaxial portion of the carpel wall which in turn overgrows the ovule and ultimately forms a long thin style with either a funnel-shaped (Vleisia, L. cylindrocarpa) , peltate (L. bilocularis) or feathery (Althenia) stigma. Relationships with other Alismatales are discussed.     

Variability of Flavonol Contents during Floral Morphogenesis in the Poppy Papaver somniferum L.

We studied the contents of flavonols (kaempferol and quercetin) in the meristem of vegetative and generative apices of the main plant shoot in floral Papaver somniferum L. mutants, as well as in the normal plants at successive stages of flower development. Five stages of flower development were distinguished. Flavonols (kaempferol and quercetin) were present in all flower organs at all stages of floral morphogenesis we studied. However, their contents and distribution in different organs and at different stages of flower development markedly varied. No significant differences were found in the contents of flavonols in the meristems of vegetative and generative apices of the main shoot in the lines of floral mutants, as well as between the lines with different amounts of vegetative phytomeres. In the plants with normal flower structure, the contents of flavonols (kaempferol + quercetin) sharply increased with the beginning of differentiation of flower organs, i.e. from stage 3, to reach a maximum in the open flower, when gametogenesis is terminated and fertilization takes place. The level of flavonol contents in the petals (upper part) and stamen was at a maximum at all stages of flower development, while that in the gynaecium was at a minimum. The kaempferol : quercetin ratio was shifted towards quercetin at successive stages of flower development, most significantly in the stamens. The involvement of flavonols in the regulation of floral morphogenesis at stages of flower organs differentiation and functioning is discussed.     

盐胁迫对苹果器官中钙镁铁锌含量的影响

以盆栽2年生富士苹果树(砧木为平邑甜茶M.hupehensisReld)为试材,研究了盐胁迫对苹果矿质营养平衡的影响.结果表明,在盐胁迫下,苹果各器官不同时期的单位干样中Ca、Mg、Fe和Zn含量的平均值没有明显变化,但各元素与Na的比值明显下降,特别是在高盐(3‰NaCl)胁迫下下降更为明显,从而破坏了树体内元素平衡.在无盐和盐胁迫下,苹果各器官中Ca含量的顺序为主干韧皮部＞叶片、新梢＞根＞主干木质部;Mg含量为新梢、根＞主干木质部、主干韧皮部、叶片;Fe含量为根＞叶片＞主干韧皮部、新梢＞主干木质部;Zn含量为新梢＞叶片＞根、主干韧皮部＞主干木质部.与对照相比,器官中各元素含量在胁迫期间表现出不同程度的波动性.     

Variability of flavonol contents during floral morphogenesis in Papaver somniferum L

We studied the contents of flavonols (kaempferol and quercetin) in the meristem of vegetative and generative apices of the main plant shoot in floral Papaver somniferum mutants, as well as in the normal plants at successive stages of flower development. Five stages of flower development were distinguished. Flavonols (kaempferol and quercetin) were present in all flower organs at all stages of floral morphogenesis we studied. However, their contents and distribution in different organs and at different stages of flower development markedly varied. No significant differences were found in the contents of flavonols in the meristems of vegetative and generative apices of the main shoot in the lines of floral mutants, as well as between the lines with different amounts of vegetative phytomeres. In the plants with normal flower structure, the contents of flavonols (kaempferol + quercetin) sharply increased with the beginning of differentiation of flower organs, i.e. from stage 3, to reach a maximum in the open flower, when gametogenesis is terminated and fertilization takes place. The level of flavonol contents in the petals (upper part) and stamen was at a maximum at all stages of flower development, while that in the gynaecium was at a minimum. The kaempferol: quercetin ratio shifted towards quercetin at successive stages of flower development, most significantly in the stamens. The involvement of flavonols in the regulation of floral morphogenesis at stages of flower organs differentiation and functioning is discussed.     

Nitrogen Uptake and Plant Growth II. An Empirical Model of Vegetative Growth and Partitioning

An empirical model of vegetative plant growth is presented.The model is based on experimental data on Polygonum cuspidatum,which showed (1) that the partitioning of dry matter and nitrogenamong organs was linearly related to the nitrogen concentrationof the whole plant and (2) that leaf thickness was negativelycorrelated with leaf nitrogen concentration. The model properlydescribes the behaviour of plants. Steady-state solutions ofthe model give the relative growth rate, specific leaf weight,and partitioning of dry matter and nitrogen among organs withthe net assimilation rate and the specific absorption rate asenvironmental variables. The effect of nitrogen removal on drymatter and nitrogen partitioning was examined as non-steady-statedynamic solutions of the model. The model predicted not onlyreduced leaf growth and enhanced root growth but also a fluxof nitrogen from the leaf to the root, which agreed with theexperimental results. Mathematical model, partitioning of dry matter and nitrogen, plant nitrogen, relative growth rate, shoot: root ratio, specific leaf weight     

Morphology and development of the reproductive shoots of Lilaea scilloides (Poir.) Hauman (Alismatidae)

Shoots of adult plants of Lilaea scilloides have a sympodial form. Each unit of the sympodium bears a single sheathing prophyll (which is the only kind of foliage leaf produced in the adult) and terminates in an inflorescence. The prophyll subtends the next unit of the sympodium. A further accessory bud can form in association with each unit. This bud repeats the pattern of the main sympodium, giving the plant a tufted habit. Five different kinds of flower can be identified in the inflorescence: a unisexual male flower with a single perianth member and adnate stamen; a bisexual flower, with a single perianth member and adnate stamen, and a single carpel with an anatropous bitegmic ovule; a unisexual female flower with a single perianth member and carpel; a unisexual female flower comprising only a single carpel; and a female flower comprising only a single carpel with a very long filamentous style. The first four kinds occur in the upper part of the inflorescence which is normally elevated on a scape, while the last kind is restricted to the base of the inflorescence. In the position of the basal flowers several variations have been observed in cultivated material. These include branching associated with the basal flowers, which results in the development of additional basal flowers or inflorescences, and even total replacement of a basal flower by an inflorescence or a branching structure bearing flowers. A review of past literature includes a clarification of some persistent errors which have confused the taxonomic position of the plant and the morphological interpretation of the reproductive appendages.     

NORMAL FLORAL ONTOGENY AND COOL TEMPERATURE-INDUCED ABERRANT FLORAL DEVELOPMENT IN GLYCINE MAX (FABACEAE)

Floral onset in soybean (Glycine max cv. Ransom) is characterized by precocious initiation of axillary meristems in the axils of the most recently initiated leaf primordium. During floral transition, leaf morphology changes from trifoliolate leaf with stipules, to a three-lobed bract, to an unlobed bract. Soybean flowers initiated at 26/22 C day/night temperatures are normal, papilionaceous, and pentamerous. Sepal, petal, and stamen whorls are initiated unidirectionally from the abaxial to adaxial side of the floral apex. The median sepal is located abaxially and the median petal adaxially on the meristem. The organogeny of ‘Ransom’ flowers was found to be: sepals, petals, outer stamens plus carpel, inner stamens; or, sepals, petals, carpel, outer stamens, inner stamens. The outer stamen whorl and the carpel show possible overlap in time of initiation. Equalization of organ size occurs only within the stamen whorls. The sepals retain distinction in size, and the petals exhibit an inverse size to age relationship. The keel petals postgenitally fuse along part of their abaxial margins; their bases, however, remain free. Soybean flowers initiated at cool day/night temperatures of 18/14 C exhibited abnormalities and intermediate organs in all whorls. The gynoecium consisted of one to ten carpels (usually three or four), and carpel connation varied. Fusion of keel petals was often lacking, and stamen filaments fused erratically. Multiple carpellate flowers developed into multiple pods that were separate or variously connate. Intermediate type organs had characteristics only of organs in adjacent whorls. These aberrant flowers demonstrate that the floral meristem of soybean is not fixed or limited in its developmental capabilities and that it has the potential to produce alternate morphological patterns.     

Dry Matter Partitioning in a Determinate and an Indeterminate Cultivar of Vicia faba L. Under Contrasting Plant Distributions and Densities

Models for root: shoot, vegetative: generative and stem: leafpartitioning are presented to quantify dry matter partitioningof two contrasting genotypes of Vicia faba, an indeterminateand a ‘topless’. A third plant type, a determinateof which two to three inflorescences had been removed, was alsoincluded but behaved similarly to the intact plant. The root:shoot partitioning model predicts linear relationships betweenroot proportion and the product of air vapour pressure deficitand relative growth rate. Data from field experiments were consistentwith model prediction and coefficients estimated were similarfor both genotypes. Partitioning into pods was modelled as proportionalto the number of actively growing pods younger than 1000°Cd. Coefficients estimated were similar in both genotypes butdifferent between densities and years. In the indeterminategenotype, stem: leaf ratio was allometric throughout, whereasleaf growth ceased but stem growth continued in the determinateafter formation of the terminal inflorescence. Relatively moredry matter was allocated to stems than to leaves in high thanin low densities. In conclusion, the main differences in drymatter partitioning between genotypes concern leaf: stem partitioningduring early pod filling and pod partitioning due to pod numberdifferences. Partitioning, root, shoot, leaf, stem, Vicia faba L.     

Changes in polyamine pattern mediates sex differentiation and unisexual flower development in monoecious cucumber (Cucumis sativus L.)

Changes in the levels of polyamines are associated with fundamental physiological processes such as embryogenesis, induction of flowering, fruit development and ripening, senescence, and responses to environmental stresses, but the role of polyamines in sex differentiation and unisexual flower development has not been deeply studied. To extend the knowledge on the regulatory mechanisms of flowering in monoecious plant (producing unisexual flowers), we investigated the morphogenesis and free polyamine levels in Cucumis sativus during sex differentiation and unisexual flower development in vitro using histocytological and biochemical methods. As shown in our study, floral development in vitro was undisturbed and flowers of both sexes were produced. Sex differentiation relied on preventing the development of generative organs of the opposite sex, as we observed carpel repression in male flowers and stamen repression in female flowers. Pollen viability was negatively correlated with female flower development on the same node. Biochemical analysis revealed increased accumulation of aliphatic amines (tri, tetra‐amines) in generative (flower buds and flowers) compare to vegetative (axillary buds and leaves) organs. Undifferentiated floral buds contained elevated levels of agmatine, cadaverine, spermidine and spermine. Sex differentiation was associated with significantly decreased levels of agmatine and cadaverine. Our results showed that female flowers contained higher levels of total polyamine than male flowers. The increased level of cadaverine was associated with macrogametogenesis and female flower maturation. Putrescine was important for male flower development. Such results support the hypothesis that aliphatic amines are involved in unisexual flower development.     

The effect of partial flower removal on the relation betwen root, shootand fruti growth in the indeterminate tomato

The root, vegetative shoot and fruit growth of November and January sown glasshouse tomato plants grown in flowing water culture was followed over 6–7 months. The relationship between vegetative and reproductive growth was examined after two-thirds of the flowers were removed from half the experimental plants. This resulted in larger plants which had fewer, larger fruits and eventually a fruit yield almost as large as the controls. In the control plants, fruit growth increased steadily until it reached 90% of the total incremental fresh weight of the plant 50–60 days after first anthesis. Leaf growth was markedly depressed at this stage and root growth ceased 4 wk after anthesis. Some root death was observed from anthesis onwards. When fruit growth subsequently diminished, vegetative growth recovered but to a lower rate than before fruiting commenced. Following partial flower removal, only 64% or less of the total increment of fresh weight went into the fruit. Although vegetative growth at this stage was thus greater than in the control plants, both shoot and root growth followed the same qualitative pattern with time. The ratio of vegetative shoot to root fresh weight remained essentially constant throughout the fruiting phase in plants of both sowings whether flowers were removed or not. This suggests that the fruit grew in competition with the vegetative organs as a whole, although, for a short period at early fruiting, root growth was more seriously affected. The pattern and amount of fruiting in this indeterminate plant was influenced by the size of the vegetative organs at fruiting, and by the effect of the existing developing fruit on further vegetative and reproductive growth.     

Organ-specific analysis of phenylphenalenone-related compounds in Xiphidium caeruleum

The distribution pattern of phenylphenalenone-type compounds was investigated in vegetative and reproductive organs of Xiphidium caeruleum. The highest total molar concentration, up to 30 micromol g(-1) fr. wt, was detected in the root tip and the stamen. Accumulation of specific phenylphenalenone-related metabolites including glycosides was found in the hypogeal plant parts, the leaves, and the reproductive organs of the inflorescence. Putative biosynthetic relationships and the role of these compounds in plant defence are discussed.     

黄土高原优势灌丛营养器官化学计量特征的环境分异和机制

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。     

Male and female flowers of the dioecious plant sorrel show different patterns of MADS box gene expression.

Male and female flowers of the dioecious plant sorrel (Rumex acetosa) each produce three whorls of developed floral organs: two similar whorls of three perianth segments and either six stamens (in the male) or a gynoecium consisting of a fertile carpel and two sterile carpels (in the female). In the developing male flower, there is no significant proliferation of cells in the center of the flower, in the position normally occupied by the carpels of a hermaphrodite plant. In the female flower, small stamen primordia are formed. To determine whether the organ differences are associated with differences in the expression of organ identity genes, cDNA clones representing the putative homologs of B and C function MADS box genes were isolated and used in an in situ hybridization analysis. The expression of RAD1 and RAD2 (two different DEFICIENS homologs) in males and females was confined to the stamen whorl; the lack of expression in the second, inner perianth whorl correlated with the sepaloid nature of the inner whorl of perianth segments. Expression of RAP1 (a PLENA homolog) occurred in the carpel and stamen whorls in very young flower primordia from both males and females. However, as soon as the inappropriate set of organs ceased to develop, RAP1 expression became undetectable in those organs. The absence of expression of RAP1 may be the cause of the arrest in organ development or may be a consequence.     

The Control of Runner Development in the Strawberry Fragaria ananassa Duch

Factors controlling runnering of strawberry plants were studiedunder controlled and semi-controlled environments. Increasinglylong exposures to conditions favourable for runnering causeda subsequent increase in number of runners, their total length,number of daughters, and of dry matter produced by mother plants.Leaf and crown numbers of the mother plant showed no such parallelincrease. Temperature supplement to foliage, combined with alight interruption in the dark period, produced the greatestvegetative effect-except again on leaf number-when comparedto either factor individually, normal day extension for threehours by itself or combined with temperature supplement. Itwas suggested that the promotive effects of temperature andphotoperiod on runnering were due to increased activation ofvegetative buds on the rosette crown. Upon comparing constantand diurnally fluctuating temperatures under a constant longphoto-period in the phytotron it was found that, while all othermeasured vegetative criteria were closely similar, the totalrunner length produced by plants under fluctuating temperatureswas almost double that of those under constant temperatures.A temperature gradient from root to shoot stimulated vegetativegrowth in comparison to a gradient in the opposite directionand to lack of such gradient, irrespective of whether the temperaturecommon to both root and shoot was high or low. This was interpretedas resulting from either different temperature requirementsfor aerial and subterranean organs or from the occurrence oftranslocation gradients of substances which promote vegetativedevelopment of the shoots, e.g. kinins and gibberellin-likesubstances. The time interval between rooting of daughter plants and onsetof runnering was increased the later the rooting occurred, possiblydue to the advancing seasonal decline in conditions favourablefor vegetative development. The positional influence of theorder of the daughter along the runner chain, which was markedin flowering, was not found.     

Auxin Depletion from the Leaf Axil Conditions Competence for Axillary Meristem Formation in Arabidopsis and Tomato

The enormous variation in architecture of flowering plants is based to a large extent on their ability to form new axes of growth throughout their life span. Secondary growth is initiated from groups of pluripotent cells, called meristems, which are established in the axils of leaves. Such meristems form lateral organs and develop into a side shoot or a flower, depending on the developmental status of the plant and environmental conditions. The phytohormone auxin is well known to play an important role in inhibiting the outgrowth of axillary buds, a phenomenon known as apical dominance. However, the role of auxin in the process of axillary meristem formation is largely unknown. In this study, we show in the model species Arabidopsis thaliana and tomato (Solanum lycopersicum) that auxin is depleted from leaf axils during vegetative development. Disruption of polar auxin transport compromises auxin depletion from the leaf axil and axillary meristem initiation. Ectopic auxin biosynthesis in leaf axils interferes with axillary meristem formation, whereas repression of auxin signaling in polar auxin transport mutants can largely rescue their branching defects. These results strongly suggest that depletion of auxin from leaf axils is a prerequisite for axillary meristem formation during vegetative development.     

NaCl处理对空心莲子草营养器官解剖结构的影响

利用光学显微技术,对不同浓度NaCl处理后的空心莲子草的营养器官进行形态解剖学研究。结果表明:不同浓度的NaCl处理下生长的空心莲子草解剖结构与对照相比发生了显著变化,这些变化表现为:随着盐浓度的逐渐增加,叶片面积变小而厚度增加;茎的横向生长受抑制;角质膜进一步增厚;营养器官的通气组织进一步发达,根、茎、叶中的胞间隙或孔隙紧密相连,使植物体上下贯通进行气体交换;根中的木栓细胞层数增加。     

NaCl胁迫下高纬度移植桐花树幼苗的生理生态效应

通过砂培试验,研究了不同浓度NaCl(0、100、200、300和400 mmol·L-1)处理对高纬度移植桐花树幼苗生物量、离子吸收、碳氮代谢、叶片光合色素和叶片抗氧化系统的影响.结果表明:100 mmol·L-1NaCl处理对桐花树生长有轻微的促进作用,当NaCl浓度达到300mmol·L-1时,桐花树根、茎、叶器官的干鲜质量、根冠比、株高和基径均显著下降.高盐胁迫抑制了叶片抗氧化系统中超氧化物歧化酶(SOD)和过氧化物酶(POD)的活性,提高了丙二醛含量,降低了叶片中叶绿素、类胡萝卜素总量以及根茎叶的可溶性总糖和游离氨基酸总量.不同浓度NaCl胁迫下,桐花树根茎叶中Na+含量快速上升,K+含量相对下降,K+/Na+快速下降,导致各器官中离子平衡失调.当NaCl浓度高于300 mmol·L-1时,桐花树根茎叶的碳氮代谢运转失调,抑制了植株的正常生长,导致各器官的生物量显著下降.     

Ectopic expression of Kxhkn5 in the viviparous species Kalanchoe?��?Houghtonii induces a novel pattern of epiphyll development

KxhKN5 (class 1 KNOX gene) was cloned from Kalanchoe × houghtonii with strong tendency to form epiphylls on leaves. KxhKN5 appear to be homologue of BP of A. thaliana on the basis of phylogeny, expression and phenotype analysis. Beside the modification of several plant and leaf traits, the appearance of epiphylls was drastically reduced by both the silencing and the over-expression of KxhKN5 in most of the generated clones. In silenced clones, epiphyll production followed the morphogenetic pathway of the WT plants: somatic embryos outbreak in the centre of each leaf-pedestal, grown in the notch between leaf indentations and were supported by a suspensor. The connection between the epiphyll and the mother plant did not include any vasculature and as a result, the epiphylls dropped easily from the mother plant. The most represented category of over expressor clones, disclosed a novel pattern of epiphyll development: the leaf-pedestals were absent and single bud outbreaks in each leaf notch. Buds developed into shoots which remained attached to the maternal plant by a strong vascular connection. The leaves supporting shoots, produced a thickened midrib and veins, and their lamina ceased expanding. Finally, the leaf/shoot structure resembles a lateral branch. The ectopic expression of KxhKN5 in K. × houghtonii induces a process comparable to the alternation of leaf and shoot formation in other species and support the idea, that it is the variation in shared molecular and developmental processes which produces the growth of specific structures.