LEAF DEVELOPMENT AND PRIMARY VASCULAR ORGANIZATION IN SHOOTS OF ANISOPHYLLEA DISTICH A

Anisophyllea disticha is characterized by strong shoot dimorphism. Orthotropic shoots with helically arranged scale leaves produce tiers of plagiotropic shoots, while plagiotropic shoots are anisophyllous and bear dorsal scale and ventral foliage leaves arranged in a unique tetrastichous system. In this study we compare the patterns of leaf development and primary vascular organization in the two types of shoots. Orthotropic shoots have an open vascular system with five sympodia. Expansion of orthotropic shoot scale leaves occurs from P1 to P10–12, and leaf tissues mature precociously. Plagiotropic shoots have a closed vascular system with six sympodia. Leaves in ventral and dorsal orthostichies do not differ significantly in size until ca. P15, but ventral leaves are distinct histologically from the second node in an orthostichy, P4–6. Ventral foliage leaves have a diffuse plate meristem, and leaf expansion continues until ca. P30. Differentiation of ventral and dorsal leaf trace procambium parallels the divergent patterns of leaf expansion. These observations demonstrate the strong correlation among shoot symmetry, leaf development, and vascular differentiation within dimorphic shoots of one species.     

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.     

HETEROPHYLLIC DEVELOPMENT IN MUEHLENBECKIA (POLYGONACEAE)

Flowering shoots of Muehlenbeckia platyclados Meisn. bear only reduced scale leaves which resemble the membranous sheath portion (ochrea) of leaves of other members of the Polygonaceae. Shoots propagated from cuttings bear enlarged foliage leaves with distinct lamina, petiole, and ochrea zones. The developmental basis for this heterophylly is explored in order to determine whether scale leaves resemble foliage leaves in their pattern of ontogeny or are developmentally unique. SEM and histological analyses have shown that scale leaves and foliage leaves are distinctive from inception. The scale leaf arises as a collarlike growth and extends over the shoot apex as a hooded sheath without evidence of blade initiation. By contrast, the first stage of foliage-leaf ontogeny is the differentiation of the distal lamina from the future leaf base. As the foliage-leaf ochrea encircles the stem axis, the lamina grows erect and projects from the abaxial surface of the sheath. Lamina reduction coupled with ochrea elaboration in intermediate leaf types indicate a homology between the entire scale leaf and foliage-leaf ochrea. Despite this homology, the production of the bladeless scale leaf does not involve a mere suppression of the foliage-leaf lamina. Erect growth of the saccate ochrea of the foliage leaf contrasts with the hooded expansion of the scale. Early histological differences, including contrasting rates of cell differentiation, also distinguish the two organs. This disparity in modes of growth and differentiation from inception results from separate, predetermined courses of ontogeny. Unlike other plants studied, leaf size and degree of leaf elaboration decrease with shoot meristem enlargement in Muehlenbeckia. Leaf packing does increase with shoot development and may contribute to variations in leaf morphology. It is concluded that the peculiarities of the heterophyllic leaf sequence in Muehlenbeckia are a property of the shoot system as a whole.     

Structure of grafted crown-gall teratoma shoots of tobacco: Regulation of transformed cells

Crown-gall teratomas are tumors of higher plants with an intrinsic capacity for organogenesis. The growth pattern of tobacco (Nicotiana tabacum L.) teratoma shoots, which is highly aberrant in primary tumors, becomes normal when the shoots are grafted to healthy stock plants. However, certain abnormalities commonly persist; tumors form at the graft junctions, leaves are small, apical dominance is incomplete, the stem and proximal region of the leaf midribs swell excessively, and localized eruptions of neoplastic growth occur on the swollen tissue. Swelling of the shoots is primarily the result of cell hypertrophy in the cortex. Neoplastic divisions do not occur as a general rule; they are restricted, with the exception of tumor formation at the graft junctions, to localized eruptions of teratoid growth on the nodes and leaf midribs where cell hypertrophy is most evident. The histology of the apical meristem and histogenesis of primary tissues is normal, even in grossly distorted shoots. Similarly, there is no evidence of unregulated division in the vascular cambium. It is concluded that cell expansion and division are tightly regulated in meristematic regions of teratoma shoots whereas post-meristematic tissue is prone to excessive hypertrophy and eventual initiation of neoplastic cell division.     

Growth of the Stem of Agropyron repens (L.) Beauv

The growth rate of the stem of Agropyron repens (L.) Beauv.begins to decline when the sixth foliage leaf has expanded butthe relative growth rate declines throughout the period betweenthe production of one and ten mature leaves. On an absolutetime scale there is a progressive decline in growth rate ofsuccessively formed stem (node-internode) units. On a plastochronscale the relative growth rate of successive stem units declineswithin the apical region but increases behind the apex. Thedecline in the apical region is related to a decrease in therate of cell division and in the later formed stem units thereis no significant increase in cell number from the time of theirformation by the apex until the internode is initiated duringtheir fourth plastochron. These changes are related to concurrentchanges in the size of the shoot apex and in rates of leaf growth.     

The Differentiation of the Leaf Cells of Wheat and the Development of Its Chloroplasts in the Mesophyll Cells

1. By means of cell separation method, we studied the differentiation of the leaf cells of wheat, Nongda 183 and the development of the chloroplasts in the mesophyll. cells. 2. The differentiation of the cells of the first leaf can be divided into 3 stages. Beginning from the leaf primordium to the fully expanded leaf, the cells are in the stage of division and expansion. When the fully expanded leaf becomes deep green in color, the leaf cells are in the prime of life. When the leaf begins to show yellowish colored spots to its complete withering, the cells are in the stage of senescence. Accompanying these stages, the external form and the internal structure of the cells change also. 3. In the early stage of cell division and expansion, one can observe many 0.5μ × 3.4μ mitochondria-like protoplastids which go through various morphological changes to become chloroplasts. 4. The mesophyll cells of the leaf begin to show the signs of senescence sooner than the epidermal cells and the cells of the vascular bundle. The latter last the longest in the life span of the leaf.     

Transverse Vein Differentiation Associated with Gas Space Formation--Fate of the Middle Cell Layer in Leaf Sheath Development of Rice

In monocotyledons, the leaf vascular network consists of a hierarchicalsequence of vertical vascular bundles and numerous transverseveins that interconnect adjacent vertical veins. In the leafsheath of these species, especially grasses, lysigenous gascavities (gas spaces) are developed into intervascular spacesand provide a gas conducting system to non-aerial parts underflooded conditions. The spatial relationship between gas spaceformation and transverse vein differentiation was investigatedusing the leaf sheath of rice (Oryza sativa L.). Histochemicalobservation showed that patterns of differentiation of the transversevein are distinct from those of vertical vascular bundles. Onthe other hand, gas spaces are formed through the processesof cell death (collapse). Both events are initiated at a specificcell position in the middle layers of the leaf sheath, fromwhich the vascular system of the leaf is derived; this indicatesthat differentiation of transverse veins is associated withgas space formation. The cell-to-cell movement of fluoresceinisothiocyanate-conjugated dextran injected into middle layercells coincided with the area where cell collapse occurred,indicating a close relationship between the middle and adaxialcell layers, but not abaxial cell layers. A uniform cell numberbetween each transverse vein in the leaf sheath suggested theinvolvement of spatial regulation in transverse vein formationregardless of clonal history at the later stage of leaf veincanalization. Copyright 2000 Annals of Botany Company Cell collapse, leaf development, middle cell layer, microinjection, Oryza sativa L., rice, programmed cell death.     

The relative share of graminoid and forb life-forms in a natural gradient of herb layer productivity

The proportional share of graminoid and forb life-form in the herbaceous layer was investigated along a productivity gradient at Laelatu, western Estonia With an increase in the herbaceous layer standing crop from 43 5 to 723 7 g m⁻² the graminoid life-form became dominant in total above-ground mass and in species number Three hypotheses to better explain competitive ability of graminoids were tested 1) graminoids are able to form higher foliage, 2) they are able to distribute foliage nitrogen in a more beneficial way, 3) they have better nitrogen use efficiency 21 sample plots 50 × 50 cm were harvested All above-ground parts of vascular plants were removed by two canopy layers Vertical separation of layers were made according to the height of half light interception A species list was compiled, total and leaf masses and leaf nitrogen content of both life-forms were measured by layer ANOVA showed that there were no significant differences in vertical distribution of foliage or foliage nitrogen between life-forms in the productivity gradient, and hypotheses 1) and 2) are not supported by our data-set Hypothesis 3) is approved partly as the nitrogen concentration in graminoid foliage was 20% less than in forbs If one supposes that nitrogen retention time is equal in both life-forms then graminoids must have higher nitrogen use efficiency when compared to forbs Although the influence of life-form x productivity interaction on leaf nitrogen concentration was not significant, there was a tendency that difference in leaf mass to nitrogen ratio of the two life-forms increased with increasing incident light Thus, we can hypothesize that graminoid species dominate in high productive plots where the incident light intensity is also higher due to their better nitrogen use efficiency when compared to forb species     

Protein Composition in Relation to Age of Daffodil Leaves

Daffodil foliage leaves were divided into sections along theirlength; the basal sections then contained the youngest, growingregions of the leaves, and the other sections represented progressivelyolder tissue as the leaf apex was approached. Representativeprotein fractions were isolated from some of these sections,and after hydrolysis their amino-acid compositions were compared.Protein from bulb scale leaves was also analysed. Within thefoliage leaf, age did not markedly affect the composition ofthe proteins. Larger differences of composition were found whenthe proteins of the bulb scale, a typical storage tissue, werecompared with those of the foliage leaves. The free amino-acid complements of the different sections ofthe foliage leaves were also compared. Variation of compositionwith leaf age did occur, but no generalizations can be madethat are applicable to all amino-acids.     

AXILLARY BUD DEVELOPMENT IN POPULUS DELTOIDES. II. LATE ONTOGENY AND VASCULARIZATION

A nearly mature axillary bud of Populus deltoides was embedded in epoxy and serially sectioned at 6 μm. Sectioning extended from the cataphyll tips to a level in the subtending internode about 6 mm below the bud base. Vascular development was followed through the serial microsections and the vascular system was mapped in its entirety from initiation of the original bud traces to termination of the last recognizable leaf trace beneath the bud apex. Each vascular trace was identified as to its origin, its termination within a foliar organ, and its relation to other traces comprising the bud vascular cylinder. Analysis of these data confirmed the procambial patterns found in Part I of this study. Two original bud traces that diverged from the central trace of the axillant leaf gave rise to two pairs of scale traces in quick succession, and these scale traces become the progenitors of all subsequent vascular traces that were perpetuated within the bud. Just before the bud vascular system separated from that of the stem, a third pair of scale traces diverged from the original bud traces; the latter then receded toward the stem to eventually merge with its vasculature. The third pair of scale traces produced a horizontal vascular connection between stem and bud before terminating in the adaxial cataphyll. The vascular system at first conformed with a ½ vascular phyllotaxy when the original bud traces were initiated, progressed through a ⅓ vascular phyllotaxy in the scale trace system, and terminated at the time of sampling with a ⅖ vascular phyllotaxy in the foliage leaf primordia.     

Growth and Development of the Banana Plant: I. The Growing Regions of the Vegetative Shoot

This is a study of the vegetative growth of the banana plant,with special reference to the structure of the shoot apex, theorigin of the leaf primordia and buds, and the growth of theleaf base into the pseudostem. The various regions in whichintercalary growth contributes to the vegetative plant bodyare described. The anatomical structures observed are illustratedby photomicrographs. Binucleate cells are conspicuous in theleaf bases and in cells produced by intercalary men-stems. Theformation of the air chambers which are characteristic of themature leaf and of the septa, which are formed as persistentsheets of cells which bound these chambers, is described. Thecell divisions which build the septa, and also those which causethe eccentric growth of the midrib are noted, and their proximityto adjacent vascular strands is stressed. Other marginal meristemsbuild the lamina of the leaf. The function of the central apicalmeristem of the shoot is not to create a massive axis whichgrows in length, for this vegetative function is taken overby the lateral organs, the growth of which greatly overshadowsthat in the main axis. However, as the vegetative shoot growsolder, its central mass of meristem does become progressivelylarger. Cell divisions in this central area are sparse, thoughsufficient to increase its bulk slowly, while the main organ-buildingand cell-multiplying functions are delegated to the lateralorgans. This condition changes on flowering when a massive,true, erect stem forms. Axillary buds do not occur in the vegetativeshoot, but adventitious buds appear in an anomalous situation.The vegetative shoot behaves as though there is an extremelystrong apical dominance, which suppresses all buds and growthin the axis itself. But an elusive question is the mechanismwhich stimulates, or controls, the behaviour of so many dividingcells, distributed so widely, through so many discrete areasof cell division or intercalary meristematic activity. The frequentproximity of vascular strands, as probable sources of both nutrientsand stimuli to cell division, is suggestive here.     

The Effect of Rooting Space on Whole Plant and Leaf Growth in Brussels Sprouts (Brassica oleracea var. gemmifera)

Seedlings of Brassica oleracea var. gemmifera DC. (Brusselssprouts) were grown in four pot sizes over a 4-week period.Whole plant, stem, root and foliage d. wts and foliage area,together with specific leaf area, leaf area ratio and numberof leaves initiated were reduced by restricting rooting space.Individual leaves showed similar reductions in d. wt and area,with the effect being more pronounced in later-formed leaves.Cell counts and measurements on the epidermis and palisade mesophylllayers of the first four leaves showed that the reduction ingrowth was due to reduced cell division. Cell numbers in thefirst-formed leaf were halved over the range of pot sizes used,and there was a progressively greater reduction in cell numbersin later-formed leaves. There was some tendency for cell sizeto decrease with decreasing rooting space, but this was notgeneral and was most marked between plants grown in the twosmallest pot sizes. Brassica oleracea var. gemmifera, Brussels sprouts, rooting space, growth analysis, leaf growth, cell numbers, cell sizes     

A review of light interception in plant stands from leaf to canopy in different plant functional types and in species with varying shade tolerance

Changes in the efficiency of light interception and in the costs for light harvesting along the light gradients from the top of the plant canopy to the bottom are the major means by which efficient light harvesting is achieved in ecosystems. In the current review analysis, leaf, shoot and canopy level determinants of plant light harvesting, the light-driven plasticity in key traits altering light harvesting, and variations among different plant functional types and between species of different shade tolerance are analyzed. In addition, plant age- and size-dependent alterations in light harvesting efficiency are also examined. At the leaf level, the variations in light harvesting are driven by alterations in leaf chlorophyll content modifies the fraction of incident light harvested by given leaf area, and in leaf dry mass per unit area (M _A) that determines the amount of leaf area formed with certain fraction of plant biomass in the leaves. In needle-leaved species with complex foliage cross-section, the degree of foliage surface exposure also depends on the leaf total-to-projected surface area ratio. At the shoot scale, foliage inclination angle distribution and foliage spatial aggregation are the major determinants of light harvesting, while at the canopy scale, branching frequency, foliage distribution and biomass allocation to leaves (F _L) modify light harvesting significantly. F _L decreases with increasing plant size from herbs to shrubs to trees due to progressively larger support costs in plant functional types with greater stature. Among trees, F _L and stand leaf area index scale positively with foliage longevity. Plant traits altering light harvesting have a large potential to adjust to light availability. Chlorophyll per mass increases, while M _A, foliage inclination from the horizontal and degree of spatial aggregation decrease with decreasing light availability. In addition, branching frequency decreases and canopies become flatter in lower light. All these plastic modifications greatly enhance light harvesting in low light. Species with greater shade tolerance typically form a more extensive canopy by having lower M _A in deciduous species and enhanced leaf longevity in evergreens. In addition, young plants of shade tolerators commonly have less strongly aggregated foliage and flatter canopies, while in adult plants partly exposed to high light, higher shade tolerance of foliage allows the shade tolerators to maintain more leaf layers, resulting in extended crowns. Within a given plant functional type, increases in plant age and size result in increases in M _A, reductions in F _L and increases in foliage aggregation, thereby reducing plant leaf area index and the efficiency of light harvesting. Such dynamic modifications in plant light harvesting play a key role in stand development and productivity. Overall, the current review analysis demonstrates that a suite of chemical and architectural traits at various scales and their plasticity drive plant light harvesting efficiency. Enhanced light harvesting can be achieved by various combinations of traits, and these suites of traits vary during plant ontogeny.     

Leaf maximum photosynthetic rate and venation are linked by hydraulics

Leaf veins are almost ubiquitous across the range of terrestrial plant diversity, yet their influence on leaf photosynthetic performance remains uncertain. We show here that specific physical attributes of the vascular plumbing network are key limiters of the hydraulic and photosynthetic proficiency of any leaf. Following the logic that leaf veins evolved to bypass inefficient water transport through living mesophyll tissue, we examined the hydraulic pathway beyond the distal ends of the vein system as a possible limiter of water transport in leaves. We tested a mechanistic hypothesis that the length of this final traverse, as water moves from veins across the mesophyll to where it evaporates from the leaf, governs the hydraulic efficiency and photosynthetic carbon assimilation of any leaf. Sampling 43 species across the breadth of plant diversity from mosses to flowering plants, we found that the post-vein traverse as determined by characters such as vein density, leaf thickness, and cell shape, was strongly correlated with the hydraulic conductivity and maximum photosynthetic rate of foliage. The shape of this correlation provided clear support for the a priori hypothesis that vein positioning limits photosynthesis via its influence on leaf hydraulic efficiency.     

Inverted vascular bundles in the leaf of Cladium (Cyperaceae)

Illustrations of seedling development and the adult plant of Cladium jamaicense Crantz are given. The course of vascular bundles in the leaf is described in detail. The structure of seedling and adult leaves are compared. Development of leaf primordia and initiation of vascular bundles are followed. Changes in the anatomy within one leaf and differences between foliage leaves and scales are related to their development. The adaxial vascular bundles that are inversely orientated are initiated within a procambial complex in close association with the largest abaxial bundles. Normally orientated adaxial bundles have an origin independent of other bundles. A hypothesis is presented which accounts for the differentiation of inverted bundles in morphogenetic terms. No developmental evidence was found to support the phylogenetic derivation of the leaf of Cladium by adaxial folding and fusion of laminar halves with their own adaxial surfaces.     

Photosynthesis and resource distribution through plant canopies

Plant canopies are characterized by dramatic gradients of light between canopy top and bottom, and interactions between light, temperature and water vapour deficits. This review summarizes current knowledge of potentials and limitations of acclimation of foliage photosynthetic capacity (A(max)) and light-harvesting efficiency to complex environmental gradients within the canopies. Acclimation of A(max) to high light availability involves accumulation of rate-limiting photosynthetic proteins per unit leaf area as the result of increases in leaf thickness in broad-leaved species and volume: total area ratio and mesophyll thickness in species with complex geometry of leaf cross-section. Enhancement of light-harvesting efficiency in low light occurs through increased chlorophyll production per unit dry mass, greater leaf area per unit dry mass investment in leaves and shoot architectural modifications that improve leaf exposure and reduce within-shoot shading. All these acclimation responses vary among species, resulting in species-specific use efficiencies of low and high light. In fast-growing canopies and in evergreen species, where foliage developed and acclimated to a certain light environment becomes shaded by newly developing foliage, leaf senescence, age-dependent changes in cell wall characteristics and limited foliage re-acclimation capacity can constrain adjustment of older leaves to modified light availabilities. The review further demonstrates that leaves in different canopy positions respond differently to dynamic fluctuations in light availability and to multiple environmental stresses. Foliage acclimated to high irradiance respond more plastically to rapid changes in leaf light environment, and is more resistant to co-occurring heat and water stress. However, in higher light, co-occurring stresses can more strongly curb the efficiency of foliage photosynthetic machinery through reductions in internal diffusion conductance to CO(2). This review demonstrates strong foliage potential for acclimation to within-canopy environmental gradients, but also highlights complex constraints on acclimation and foliage functioning resulting from light x foliage age interactions, multiple environmental stresses, dynamic light fluctuations and species-specific leaf and shoot structural constraints.     

EARLY LEAF ONTOGENY AND SHOOT DEVELOPMENT IN BROWNEA ARIZA (LEGUMINOSAE)

Brownea ariza Benth. (Leguminosae: Caesalpinioideae) shows early shoot tip abortion and subsequent renewal growth from the pseudoterminal bud. This species is unusual in that the entire shoot system is formed before flushing from the bud occurs, shoot tip abortion occurs during flushing, and the aborting portion contains three to six leaves as well as primordial structures varying from hood to peg shape. This study focused on the morphological changes from initiation of scale and foliage leaf primordia in the “resting” renewal bud through bud elongation to flushing and bud abortion. Scanning electron microscopy revealed that embryonic scale leaves are hood-shaped while foliage leaf primordia show early segmentation into leaflets and stipules. No transitional stages were observed. Bud scales and foliage leaves show opposite developmental trends. In bud scales, length at maturity increases from first to last formed, while length decreases in sequentially formed foliage leaves. Early in leaf development the stipules keep pace with the elongation of the rachis. When the bud reaches about one half of its final length the leaf rachis begins to exceed the lengths of its stipules. This young rachis terminates in a distinct mucro that persists until maturity at which time it abscises. Growth patterns indicate that mucro and rachis are a single developmental unit. The early abortion of a shoot tip containing several leaves cannot be easily rationalized. Previous suggestions have involved maintenance of form and ecological adaptation. We add the possibility of elimination of cell progeny encumbered by mutations. From this and other studies of this group, it is clear that at maturity leaves of different species may look alike, e.g., Hymenaea and Colophospermum are bifoliolate; Brownea, Saraca, and others are multifoliolate. However, early stages of leaf ontogeny are quite diverse and may be of systematic value, since these early differences are lost or masked by later development.     

THE DEVELOPMENT OF THE ACHENE OF POLYGONUM PENSYLVANICUM: EMBRYO,ENDOSPERM, AND PERICARP

A study was made of the ontogeny of the achene of Polygonum pensylvanicum L. from fertilization to maturity. The proembryo is classified as the Polygonum Variation, Asterad Type. Cotyledons are initiated three days after anthesis, and by the fifth day procambium is present in the embryo axis. At approximately seven days after anthesis, the embryo begins to curve and occupy a marginal position in the ovary. By ten days the first foliage leaf primordium is initiated at the stem apex of the embryo. At maturity the embryo consists of two cotyledons, a plumule composed of the stem apex and one leaf primordium, and a hypocotyl with a well-developed radicle. Endosperm nuclei begin to divide before the first division of the zygote. Cell wall formation begins in the endosperm at the micropylar end of the embryo sac and proceeds toward the chalazal region. By the fifth day the endosperm is completely cellular, except for a basal projection; and a peripheral meristem has been established. At approximately ten days the peripheral meristem ceases periclinal cell division and becomes the aleurone. At the time of fertilization the ovary wall has its full complement of cell layers. The walls of the outermost cells elongate and become convoluted. Subsequent thickening and lignification of these cell walls produce the hard epicarp of the mature achene.     

Leaf expansion and cell division are affected by reducing absorbed light before but not after the decline in cell division rate in the sunflower leaf

The effect of absorbed photosynthetic photon flux density (PPFD) on leaf expansion is a key issue for analysing the phenotypic variability between plants and for modelling feedback loops. Expansion and epidermal cell division in leaf 8 of sunflower were analysed in a series of five experiments where absorbed photosynthetic photon flux density (PPFD) was reduced either by shading or by covering part of the leaf area. These treatments were imposed at different times during leaf development. Expansion and cell division were affected by a reduction in absorbed PPFD only in the first part of leaf development, while the leaf area was less than 2% of its final value and while absolute expansion rate was slow. In contrast, it was not affected if imposed later when the leaf was visible and absolute expansion rate was at maximum. A reduction in absorbed PPFD caused the same reduction in expansion and in cell division whether it was due to a reduction in incident PPFD or to a reduction in photosynthetic leaf area, suggesting that carbon metabolism was involved. Relative expansion rate recovered to control levels when relative division rate began to decline, in all experiments and in all zones of a leaf. This was probably linked to the source–sink transition, after which the leaf had such a high priority in carbon allocation that it was largely insensitive to changes in absorbed PPFD. The final leaf area was therefore closely related to the cumulated PPFD absorbed by the plant from leaf initiation to the end of exponential cell division.     

荚果蕨胚胎发育的研究

利用石蜡切片法研究了荚果蕨（Matteuccia struthiopteris(L.) Todaro）胚胎发育过程。合子第一次分裂,分裂面垂直于原叶体纵轴且平行于颈卵器颈部;第二次分裂面平行于原叶体纵轴且垂直于颈卵器颈部;第三次分裂面同时平行于原叶体纵轴和颈卵器颈部。经多次分裂的球形胚胎,胚胎的外上和外下区域几乎同时分别发育出第一叶顶端细胞和第一根顶端细胞。随着发育的进行,它们分别斜向分裂产生第一叶原基和第一根原基。随后,第一叶原基迅速分裂,突破帽状体形成第一幼叶;而第一根原基的分裂速度稍慢,第一根发育速度稍慢于第一叶。