The prediction of leaf canopy expansion in the leek from a simple model dependent on primordial development

In an analysis of leaf development of leek plants grown in the field in 1988, successive leaves initiated, appeared (tip and ligule) and senesced at equal intervals of accumulated temperature/thermal time. These intervals corresponded to a plastochron of 92°C days and phyllochrons of 135 (tip) and 233 (ligule) °C days. The rate of appearance of ligules was exactly equal to the rate of leaf senescence, with the result that the number of fully-expanded leaves per plant remained constant at 1.4. These data, which were compatible with results from previous seasons, were used to develop a model of the interrelationships between primordium initiation at the shoot apex and subsequent events in the development of individual leaves. Primordium initiation is considered to be the primary controlling event in the life of a leaf, and the processes of tip appearance, ligule appearance and death can be predicted from knowledge of the number of primordia which have been initiated, without reference to the environment. A model of canopy expansion, based on the central role of the shoot apex, was developed using the temperature relations of primordium initiation and additional data on leaf expansion and leaf dimensions. Leaf area indices computed in this way provided a satisfactory simulation of the thermal-time course of leaf area index observed in a previous season, 1985.     

Following the initiation and development of individual leaf primordia at the level of the shoot apical meristem: the case of distichous phyllotaxis in Begonia

BACKGROUND AND AIMS: By using the technique of replicas of a developing apex it is possible to obtain a direct measure of phyllotactic parameters (plastochrone and platochronic ratio) involved in the initiation of two successive primordia at the level of the SAM. The goal of this study is to compare, in a real time setting, the value of phyllotactic parameters in distichous systems using Begonia as a case study, with the value of the same parameters in spiral phyllotactic systems. METHODS: To determine the real-time sequence of events at the level of the SAM, replicas were made of the developing apex at different intervals using previously described techniques. Impression moulds were made at 24-h intervals. The following phyllotactic parameters were measured: plastochrone, angle of divergence, plastochrone ratio and ratio between the diameter of the leaf and the apex. RESULTS: The time between the appearance of two successive leaves is 15-20 d. The average value of the plastochrone ratio (R) is 1.3, and the ratio of the leaf to the diameter of the apex (Gamma) is 2.5. The angle of divergence varies from 165 masculine to 180 masculine. The speed of advection of the primordium from the apex, varies from 0.28 to 0.37 microm d(-1). CONCLUSIONS: The speed of advection of primordia in Begonia is lower than that of Anagalis. This is not in accordance with theoretical simulations that predict the opposite. In Begonia, the plastochrone ratio does not reflect the real time of appearance of two successive primordia. The time separating the appearance of two primordia is not directly related to the distance of these two primordia from the centre of the apex but is related instead to the enlargement of leaves.     

Leaf Growth and Apex Development of Perennial Ryegrass During Winter and Spring

The relationships between rates of leaf extension, leaf appearance,and primordia production on the apex were studied in the fieldduring winter and spring in established swards of four contrastingperennial ryegrass lines (Aurora, Melle, a hybrid selectionfrom a cross between Aurora and Melle and S.24). All four linesshowed an increase in leaf extension rates which commenced whenspikelet primordia were first initiated at the apex. This wassome time after vernalization requirements had been satisfied.In early-flowering lines the stimulus to leaf growth rates occurredearlier than in late-flowering lines. Maximum leaf growth ratesoccurred about the time of double ridge formation, in the middleof the period of spikelet primordia production. The rate andduration of the period of spikelet primordia production variedbetween lines. By the time of flowering, leaf growth rates declinedto values recorded for vegetative plants in the winter. Leafappearance rates followed a similar pattern to leaf growth rates,although the increase in leaf appearance rate was less thanin primordia production or leaf growth rates. Hybrids from across between early- and late-flowering lines showed early enhancementof leaf extension rates due to early initiation of spikeletprimordia production. These high rates of leaf growth were maintainedfor longer, compared with the early-flowering line, as the durationof spikelet production was longer. This illustrates a mechanismfor combining early spring growth with lateness of floweringin ryegrass breeding programmes Leaf growth, apex development, vernalization, initiation, spikelet primordia production, flowering, thermal time, Lolium perenne, perennial ryegrass, spring growth     

Co-ordination between leaf initiation and leaf appearance in field-grown maize (Zea mays): genotypic differences in response of rates to temperature

BACKGROUND AND AIMS: In maize (Zea mays), early flowering date, which is a valuable trait for several cropping systems, is associated with the number of leaves per plant and the leaf appearance rate. Final leaf number depends upon the rate and duration of leaf initiation. The aims of this study were to analyse the genotypic variation in the response to temperature of leaf appearance rate and leaf initiation rate, and to investigate the co-ordination between these processes under field conditions. METHODS: Sixteen hybrids of different origins were grown under six contrasting environmental conditions. The number of appeared leaves was measured twice a week to estimate leaf appearance rate (leaves d(-1)). Plants were dissected at four sampling dates to determine the number of initiated leaves and estimate leaf initiation rate (leaves d(-1)). A co-ordination model was fitted between the number of initiated leaves and the number of appeared leaves. This model was validated using two independent data sets. KEY RESULTS: Significant (P < 0.05) differences were found among hybrids in the response to temperature of leaf initiation rate (plastochron) and leaf appearance rate (phyllochron). Plastochron ranged between 24.3 and 36.4 degree days (degrees Cd), with a base temperature (Tb) between 4.0 and 8.2 degrees C. Phyllochron ranged between 48.6 and 65.5 degrees Cd, with a Tb between 2.9 and 5.0 degrees C. A single co-ordination model was fitted between the two processes for all hybrids and environments (r2= 0.96, P < 0.0001), and was successfully validated (coefficient of variation < 9 %). CONCLUSIONS: This work has established the existence of genotypic variability in leaf initiation rate and leaf appearance rate in response to temperature, which is a promising result for maize breeding; and the interdependence between these processes from seedling emergence up to floral initiation.     

Field studies of leaf development and expansion in the leek (Allium porrum)

An analysis of leek leaf development and expansion was carried out over three seasons using field-grown plants of three varieties which were directly sown at different dates or transplanted from controlled conditions. In all cases, successive leaves appeared (tip visible) at equal intervals of accumulated temperature. Detailed analysis of a single sowing in 1985 showed that the regularity of leaf appearance was a consequence of the coordinated response to accumulated temperature of leaf initiation (plastochron 100°C days > 0°C) and leaf blade and sheath extension. For each successive leaf, an additional 32°C days were required between initiation and appearance to allow for the linear increase in ‘sheath’ height, giving a phyllochron of 132°C days. Direct measurement of leaf extension before and after leaf appearance, and of the length of the leaf extension zone, confirmed that the rate of leaf extension, in terms of accumulated tempeature, was constant, and independent of leaf number. However, there were differences between seasons and between varieties in the responses of leaf appearance, leaf extension and ‘sheath’ length to accumulated temperature. It was concluded that the simple ontogenetic increase in leaf dimensions, which was a feature of all the crops studied, was a consequence of the progressive increase in the duration of leaf expansion.     

Coordination between plant and apex development in Hordeum vulgare ssp. distichum

Developmental scales for cereals describe apex and plant morphology separately. In order to link crucial steps of internal and external development, in three varieties of Hordeum vulgare spp. distichum L., sown in autumn and in spring, we recorded plant, leaf and apex stage, following the scales of Zadoks, Haun, and Banerjee and Wienhues, the number of primordia, culm and spike length, and the final number of leaves and spikelets. Primordia initiation was coordinated with leaf appearance and the relative rate was constant for the initiation of productive primordia. The maximum number of primordia was achieved just before the first node became detectable, but development was completed only by those initiated before floret differentiation and internode distension started. The first spikelet was initiated when the third leaf tip became visible, and the last one when plants were at the pseudo stem erection stage and five leaves had still to appear.     

The influence of temperature on leaf development and growth in potatoes in controlled environments

Three controlled environment experiments were conducted at different temperatures to determine the relation between temperature and leaf development and growth in the potato (cv. Maris Piper). Developmental stages are defined for the appearance and duration of leaf extension in the potato and comparisons made with other temperate zone crops. The rate of leaf appearance was linear over the temperature range (9–25°C) and above 25°C there was no further increase in the rate. The temperature coefficient for the rate of appearance of leaves was 0.032 leaves (degree days)^-1 using a base temperature of 0°C. The duration of extension of an individual leaf decreased with increase in temperature up to 25°C such that the thermal duration was constant at 170 degree days using a base temperature of 0°C for leaf positions 4–10 on the main stem. At higher leaf positions the thermal duration was similar or greater. The advantages and limitations of controlled environment work as a parallel to field experimentation are discussed.     

Shoot development ofAucuba japonica I. Morphological study

The shoot development ofAucuba japonica was studied morphologically. The shoot shows dichasial branching in connection with the formation of a terminal inflorescence and shows a decussate phyllotaxis even in the reproductive phase. The sequence of initiation of successive foliar appendages is very precise, hence the foliage leaf, scale leaf and bract can be compared with each other even at their stages of initiation. In the stage of proximal foliage leaf formation the shoot apex is flat, while in the stage of formation of distal foliage leaves, bud scales and proximal bracts, it becomes concave. In the stage of formation of distal bracts the apex becomes domed. Plastochron durations are relatively long in the vegetative phase in comparison with other plants, and the duration from initiation of the first pair of appendages to that of the second is about one and a half months. Both male and female inflorescences exhibit basically a thyrsoid type of monotelic synflorescence.     

A role for young leaves in vernalization of cauliflower: I. Analysis of leaf development during curd induction

Growth analysis techniques are used to test the hypothesis that chilling induces curd (flower) initiation in the cauliflower ( Brassica oleracea Botrytis L. cv. Perfection) through inhibiting leaf growth, thereby increasing the availability of growth factors to the stem apex and enabling differentiation of the curd. Effects of chilling on leaf growth and curd induction are compared in juvenile and mature, vegetative plants. Chilling at 5°C reduced dry matter accumulation in the total leaf complement by ca 60% in juvenile plants and 40% in mature plants, compared to control plants growth at 20°C. Juvenile plants showed slower rates of leaf initiation than mature plants. Leaf initiation was retarded by chilling in both plant types with the most marked effect seen in the juvenile plants. This was consistent with dry matter availability to the stem apex limiting differentiation more severely in juvenile plants than in mature plants. The rate of dry matter accumulation in existing leaves, however, was faster in juvenile plants than in mature plants at 20°C. Plants that were juvenile during chilling produced an average of 43 leaves below the curd whereas those that were mature produced 25.
Dry matter accumulation in younger leaves was more markedly inhibited by chilling than in older leaves. Chilling also reduced the rate at which enlarging leaves became positionally more remote from the stem apex. Possible roles for such leaves in regulating apical development are considered.     

Growth analysis of maize field crops under phosphorus deficiency

Biomass accumulation by crops depends both on light interception by leaves and on the efficiency with which the intercepted light is used to produce dry matter. Our aim was to identify which of these processes were affected for maize (Zea Mays L., cv Volga) field crops grown under phosphorus (P) deficiency, and assess their relative importance. In this paper, the effects of P deficiency on leaf appearance, leaf elongation rate, final individual leaf area and leaf senescence were studied. The experimental work was carried out in 1995–1977 on a long-term P fertilisation trial located on a sandy soil in the south-west of France. Three P fertilisation regimes have been applied since 1972: no-P (P0 treatment) and different rates of P fertiliser (P1.5:1.5 times the grain P export and P3:3 times the grain P export). These fertilisation regimes have led to contrasted levels of soil P supply, with the P0 treatment being limiting for growth. Very few differences were observed about leaf growth between the P1.5 and P3 treatments. Conversely, the leaf area index (LAI) was significantly reduced in the P0 treatment, especially during the first phases of the crop cycle (up to −60% between the 7- and 14-visible leaves). This effect gradually decreased over time. The lower LAI in P0 treatment was due to two main processes affecting the leaf growth. The final number of leaves per plant and leaf senescence were only slightly modified by P deficiency. Conversely, leaf appearance was delayed during the period between leaf 4 and leaf 9. The value of the phyllochron increased from 47 °C days in the P1.5 treatment to 65 °C days in the P0 treatment. Leaf elongation rates during the quasi-linear phase of leaf expansion were significantly reduced for lower leaves of P0 plants. The final size of leaves L2–L12 was reduced. On the opposite, leaf elongation duration was not greatly affected by P treatments. Before the emergence of leaf 9, the reduction of individual leaf size was the main factor responsible for the reduced LAI in the P0 treatment. After this stage, the delayed leaf appearance accounted for a great part of the reduced LAI in the P0 treatment. This revised version was published online in June 2006 with corrections to the Cover Date.     

Key phenological events in globe artichoke (Cynara cardunculus var.scolymus) development

A priority for the field vegetable grower is to be able to schedule a regular supply of product throughout the growing season. This requires a predictive framework, based on the identification of key developmental events of the crop, and an understanding of how genotypic and environmental factors interact to determine plant development. Four globe artichoke (Cynara cardunculus var. scolymus) cultivars, representing the existing phenological range, were grown in a field experiment, and a range of environmental conditions was imposed by varying both the timing of the first irrigation (which determines the initiation of regrowth) and by repeating the experiment across two locations and 2 years. The timing of the appearance of the main stem capitulum was sensitive to both the growing environment and the cultivar. These differences persisted till flowering and were correlated with final leaf number. As the plant developed, the phyllochron decreased, resulting in three values of phyllochron, each of which was responsive to genotype, and hardly to environment. The timing of the first change in phyllochron was associated with the final leaf number and the appearance of the capitulum. For all the cultivars, the rate of development fell and the final leaf number increased as the length of the photoperiod increased. The later flowering cultivars shared a similar vernalisation requirement, but ‘Spinoso sardo’ did not require a cold period to flower. Leaf length reached a peak before the beginning of stem elongation, and maximum leaf length was correlated with final leaf number. The sensitiveness of the phyllochron to the genotype, and of the number of leaves and the timing of the appearance of the capitulum to both genotype and environment makes them suitable as variables in developmental models. The importance of the final number of leaves is not only because of its phenological significance, but also because of its effect on the ability of the canopy to intercept radiation.     

The setting of rates of development of wheat plants at crop emergence: Influence of the environment on rates of leaf appearance

Examination of the time courses, expressed in terms of accumulated temperature over 0°C, of leaf appearance of fifteen sowings of winter wheat, including five cultivars, over three seasons at Auchincruive, revealed several examples of bi-linear rather than linear responses. Stepwise linear regression analysis, varying the temperature threshold by intervals of 1°C, showed that a linear relationship could be obtained for each sowing, but the appropriateness of this analysis is questioned. An alternative interpretation, that such changes in rate do have biological rather than purely statistical significance, is supported by the fact that, when they occurred, they coincided with the change in rate of spikelet initiation by the stem apex. Furthermore, similar relationships between leaf appearance and reproductive development have been demonstrated for perennial ryegrasses. Analysis of each sowing also revealed the existence of sub-populations of plants with different rates of leaf appearance and final leaf numbers, but with synchronous phenologies. These findings are discussed in relation to the coordination of the development of apex and leaves, and the environmental cues required to ensure this coordination.     

Leaf primordia initiation,leaf emergence and tillering in wheat (Triticum aestivum L.) grown under low-phosphorus conditions

In two simultaneous experiments we examined the effects of phosphorus (P) supply on leaf area development in wheat (Triticum aestivum L.) grown in sand with nutrient solutions. In Experiment 1 we studied leaf emergence, leaf elongation, tiller emergence, shoot growth, and P uptake under four levels of P supply (mM) 0.025 (P1), 0.05 (P2), 0.1 (P3), and 0.5 (P4), and. In Experiment 2 there were two levels of P supply, P1 and P4, and we examined the effects of P on leaf primordia differentiation and leaf emergence. The phyllochron was calculated as the inverse of the rate of leaf emergence calculated from the regression of number of leaf tips (PHY-Ltip), Haun index (PHY-Haun), and as the cumulated thermal time between the emergence of two consecutive leaves (PHYtt). The plastochron was calculated from the inverse of the rate of leaf primordia initiation in the apex. P deficiency delayed the emergence of leaves on the main stem and on the tiller 1. Phosphorus deficiency increased the time from emergence to double ridge and anthesis. The final number of leaves was not affected by P. The effects of P on the value of the phyllochron were attributed to both a reduced rate of leaf primordia initiation, and to a reduced leaf elongation rate. P deficiency delayed or even suppressed the emergence of certain tillers. In this work a phosphorus deficiency that reduced shoot growth by 25% at 44 days after emergence significantly modified the structure of the plants by increasing the value of the phyllochron and delaying tillering. These results suggest that any attempt to simulate leaf area development and growth of wheat plants for P-limited conditions should include the effects of the deficiency on leaf emergence.     

A phenological scale for the development of Gladiolus

A staging system for development of gladiola (Gladiolus × grandiflorus) that relies on simple, visual, non‐destructive criteria is proposed. Four field trials were conducted during the spring 2010, autumn/winter 2011 and winter 2011 at Santa Maria, RS, Brazil, with different gladiola cultivars, in order to observe the developmental stages of the above‐ground parts and their dry matter. The developmental cycle, which starts at dormant corm and ends with plant senescence, is divided into four developmental phases: dormancy phase, sprouting phase (from filiform roots appearance to sheaths appearance), vegetative phase (from emergence of the first leaf tip to emergence of the final leaf tip on the stem) and reproductive phase (from heading to plant senescence). The developmental stages that were identified during the dormancy phase and during the sprouting phases are coded as S stages: S0 = dormant corm, S1 = appearance of roots, S2.1 = first sheath, S2.2 = second sheath and S2.3 = third sheath. Vegetative phase is coded as V stages: VE = emergence of the sheaths above ground, V1 = first leaf, V2 = second leaf, Vn = nth leaf and VF = flag leaf. Leaf tip is the marker for V1–VF. The developmental stages during the reproductive phases are coded as R stages: R1 = heading, R2 = blooming, R3 = onset of flowering, R4 = end of anthesis, R5 = end of florets senescence and R6 = plant senescence (leaves and floret axis are brown). Sub‐stages have also been assigned between R1 and R2 and between R3 and R4. Illustrations (photographs) of each developmental stage taken from field pot‐grown plants are provided and the proposed scale was tested with field observations. These criteria are straight forward and allow for quick determination of development stage. This system can be used by both farmers and for experimental trials.     

Spatial and temporal analysis of non-steady elongation of rice leaves

A precise knowledge of the temporal and spatial distributions of cell division and tissue expansion is essential for appropriate leaf sampling in omics studies and for analyses of plant–environment relations. Elongating leaves of rice were studied during their whole development for elongation rate, distribution of cell length, cell production rate and spatial distribution of growth in the leaf. In seven genotypes, the pattern of leaf elongation rate followed three phases: (1) an exponential increase before leaf appearance; (2) a short phase (2–4 d at 20 °C) with a stable leaf elongation rate around leaf appearance; and (3) a phase of 8–10 d with a progressive decrease in elongation rate. The profile of cell length along the leaf changed with time during the first and last phases, but was time invariant around appearance. We propose a method adapted to non-steady elongation based on anatomical measurements, which was successfully tested by comparing it with the pricking method. It allowed analysis of the change with time in the spatial distribution of growth from initiation to end of leaf growth. The length of leaf zones with cell division and tissue elongation varied with time, with maximums of 21 and 60 mm respectively around leaf appearance.     

Effects of Shading the First Leaf on Growth of Barley Plants: I. Long-term Experiments

Plants of barley were grown under controlled conditions andthe first or second leaves covered with tubular shades thusreducing the light intensity at the leaf surface to low levels.Expansion of the shaded leaves was not prevented, but appearanceof the next leaf but one and all subsequent leaves on the mainstemwas delayed by up to 3 days. Primordia of the first four leaveswere present in the dry grain. Shade treatment delayed slightlythe initiation of the eighth and subsequent leaves and transitionto the double ridge stage at the mainstem apex. Shading the first leaf caused a temporary reduction in the rateof dry-matter increase of plants, but after 14 days the ratewas similar to that of control plants. Smaller effects werefound when the second leaf was shaded. Dry-matter productionfollowed two logarithmic phases in the period prior to awn emergence,and rates for the whole plant and for plant parts were similarfor control and shaded plants. Thus, apart from the initialperturbation, shading had no effect on growth in terms of rateof dry-weight gain. Shade treatment did not affect weight per grain or numbers ofgrain per ear, but over-all yield of grain was significantlyreduced since shading delayed the appearance of tillers andalso reduced the number of tillers bearing grain. The effectof shade was especially marked on tillers originating on primarytillers. Similar qualitative effects on tiller development werefound in an experiment on wheat.     

Bi-directional inflorescence development inArabidopsis thaliana: Acropetal initiation of flowers and basipetal initiation of paraclades

In this study we investigated Arabidopsis thaliana (L.) Heynh. inflorescence development by characterizing morphological changes at the shoot apex during the transition to flowering. Sixteen-hour photoperiods were used to synchronously induce flowering in vegetative plants grown for 30 d in non-inductive 8-h photoperiods. During the first inductive cycle, the shoot apical meristem ceased producing leaf primordia and began to produce flower primordia. The differentiation of paraclades (axillary flowering shoots), however, did not occur until after the initiation of multiple flower primordia from the shoot apical meristem. Paraclades were produced by the basipetal activation of buds from the axils of leaf primordia which had been initiated prior to photoperiodic induction. Concurrent with the activation of paraclades was the partial suppression of paraclade-associated leaf primordia, which became bract leaves. The suppression of bract-leaf primordia and the abrupt initiation of flower primordia during the first inductive photoperiod is indicative of a single phase change during the transition to flowering in photoperiodically induced Arabidopsis. Morphogenetic changes characteristic of the transition to flowering in plants grown continuously in 16-h photoperiods were qualitatively equivalent to the changes observed in plants which were photoperiodically induced after 30 d. These results suggest that Arabidopsis has only two phases of development, a vegetative phase and a reproductive phase; and that the production of flower primordia, the differentiation of paraclades from the axils of pre-existing leaf primordia and the elongation of internodes all occur during the reproductive phase.     

Changes in the levels and composition of sterols in different tissues of Lolium temulentum plants during floral development

Free, esterified and glycosylated sterols were analysed separately from the shoot apices, leaves, leaf sheaths and stems of Lolium temulentum L. (strain Ceres) plants during floral development. Short-day grown plants (50 days old) were induced to flower by exposure to a single long day. The four major sterols found by GC-MS analysis were sitosterol, cholesterol, campesterol and stigmasterol. The sterol levels in the shoot apex were much higher than those in the leaf, leaf sheath and stem. A much greater proportion of cholesterol was found in the shoot apex than in other tissues and this may reflect a specific association of cholesterol with meristematic and/or reproductive tissues.
During the inductive treatment, the sterol levels decreased in all four tissues. The major effect during early differentiation was the occurrence of transient increases in the free and esterified sterol levels in the leaf and the stem tissues. The steryl ester content peaked 24 h before the appearance of double ridges, followed by a peak in free sterol content at the double ridge stage. Similar changes could not be detected in the shoot apices. This is the first report of the sterol composition of developing shoot apices, and the results emphasize the dynamic nature of sterol metabolism during reproductive growth of L. temulentum.     

Ontogeny of the reproductive shoot apex ofClethra barbinervis,especially on the superficial view

The structure and the ontogenetic process of the reproductive shoot apex forming a terminal inflorescence ofClethra barbinervis were examined, especially concerning the superficial view of the apex. The system of contact parastichies is 2+3 in phyllotaxis in the vegetative phase, changing to 5+8 for bract arrangement in the reproductive phase. At the same time the size of the apex is conspicuously enlarged. The size of the foliage leaf primordia in the vegetative phase is larger than that of the bract primordia in the reproductive phase. The radial cell files, which are clear in the vegetative shoot apex, are not recognizable at least in the early stage of the reproductive phase. The author proposes a close correlation between the appearance of the radial cell files, as well as the construction of the apical sectors, and the sizes of the shoot apex and leaf primordia. It may be proposed also that the construction of the apical sectors is closely correlated with the phyllotaxis.