Rice morphogenesis and plant architecture: measurement, specification and the reconstruction of structural development by 3D architectural modelling

BACKGROUND AND AIMS: The morphogenesis and architecture of a rice plant, Oryza sativa, are critical factors in the yield equation, but they are not well studied because of the lack of appropriate tools for 3D measurement. The architecture of rice plants is characterized by a large number of tillers and leaves. The aims of this study were to specify rice plant architecture and to find appropriate functions to represent the 3D growth across all growth stages. METHODS: A japonica type rice, 'Namaga', was grown in pots under outdoor conditions. A 3D digitizer was used to measure the rice plant structure at intervals from the young seedling stage to maturity. The L-system formalism was applied to create '3D virtual rice' plants, incorporating models of phenological development and leaf emergence period as a function of temperature and photoperiod, which were used to determine the timing of tiller emergence. KEY RESULTS: The relationships between the nodal positions and leaf lengths, leaf angles and tiller angles were analysed and used to determine growth functions for the models. The '3D virtual rice' reproduces the structural development of isolated plants and provides a good estimation of the tillering process, and of the accumulation of leaves. CONCLUSIONS: The results indicated that the '3D virtual rice' has a possibility to demonstrate the differences in the structure and development between cultivars and under different environmental conditions. Future work, necessary to reflect both cultivar and environmental effects on the model performance, and to link with physiological models, is proposed in the discussion.     

Location Distribution Characteristics in Leaf Lateral Asymmetry of Hybrid Indica Rice

Rice leaves show lateral asymmetry. Differences in width, thickness, chlorophyll meter readings, nitrogen content, density, and yield on each asymmetrical have been reported. However, limited information is available on the distribution characteristics of the smooth side (SS) and rough side (RS) of rice leaves. The SS of rice leaves can be divided into two types according to its location in leaf: one is the smooth side appearing on the left (SSAL) of the leaf, while the smooth side of the other type appears on the right (SSAR). Through the investigation of four rice varieties, we identified that the proportion of SSAL or SSAR of flag leaves on main stems or tillers was close to 0.50 in the rice population. On the same tiller stem, SSAL and SSAR present alternately, and the SS always stretches ahead of the RS in the curled state. Among the tiller flag leaves of a single rice plant, the proportion of tiller flag leaves with SSAL or SSAR to the total number of tillers is a discrete random variable. Frequency distribution analysis of these proportions showed that the percent of frequency of some class limits was close to each other, and the ratios of percent of frequency of SSAL to SSAR were close to 1:1. To our knowledge, this is a novel finding in rice canopy morphology. According to this study, we can divide rice population into two groups in four hybrid rice varieties.     

Tolerance and compensatory response of rice to sugarcane borer (Lepidoptera: Crambidae) injury

A 3-yr field experiment was conducted to evaluate the tolerance and compensatory response of rice (Oryza sativa L.) to injury caused by sugarcane borer, Diatraea saccharalis (F.), as affected by cultivar (Cocodrie, Francis, and Jefferson), stage of crop growth during which the injury occurred (third tiller stage, panicle differentiation stage, and heading stage), and sugarcane borer density. The proportion of rice tillers with sugarcane borer injury (leaf and leaf sheath injury and/or stem injury) was lower when injury occurred at the third tiller stage (0.05) than at panicle differentiation (0.19) and heading (0.18). When injury occurred at the two latter stages, both the proportion of tillers with injury and the proportion of tillers with stem injury were negatively correlated with rainfall. Rainfall resulted in dislodgement and mortality of sugarcane borer eggs and larvae before the larvae entered the stems. Rice plant density in this study (111.1 plants/m2) was higher than recorded for previous research on rice compensation using potted rice or conducted in low-density hill production systems (26.7-51.3 plants/m2). Two mechanisms of within-plant tolerance/compensation were observed. Stem injured plants produced approximately 0.69 more tillers than uninjured plants, whereas tillers with leaf and leaf sheath injury produced larger panicles, up to 39.5 and 21.0% heavier than uninjured tillers, when injury occurred at third tiller stage and at panicle differentiation, respectively. Rice yield was not reduced with up to 23% injured tiller and up to 10% injured stems at the third tiller stage, 42% injured tillers and 17% injured stems at panicle differentiation, and 28% injured tillers and 14% injured stems at heading. Significant between-plant compensation was not detected, suggesting competition between adjacent plants is not significantly reduced by injury. Our results suggest that rice can tolerate and/or compensate for a level of stem borer injury previously considered to be economically damaging.     

Tillering in grain sorghum over a wiide range of population densities: modelling dynamics of tiller fertility

The prediction of tillering is poor or absent in existing sorghum crop models even though fertile tillers contribute significantly to grain yield. The objective of this study was to identify general quantitative relationships underpinning tiller dynamics of sorghum for a broad range of assimilate availabilities. Emergence, phenology, leaf area development and fertility of individual main culms and tillers were quantified weekly in plants grown at one of four plant densities ranging from two to 16 plants m(-2). On any given day, a tiller was considered potentially fertile (a posteriori) if its number of leaves continued to increase thereafter. The dynamics of potentially fertile tiller number per plant varied greatly with plant density, but could generally be described by three determinants, stable across plant densities: tiller emergence rate aligned with leaf ligule appearance rate; cessation of tiller emergence occurred at a stable leaf area index; and rate of decrease in potentially fertile tillers was linearly related to the ratio of realized to potential leaf area growth. Realized leaf area growth is the measured increase in leaf area, whereas potential leaf area growth is the estimated increase in leaf area if all potentially fertile tillers were to continue to develop. Procedures to predict this ratio, by estimating realized leaf area per plant from intercepted radiation and potential leaf area per plant from the number and type of developing axes, are presented. While it is suitable for modelling tiller dynamics in grain sorghum, this general framework needs to be validated by testing it in different environments and for other cultivars.     

GROWTH AND NUTRITION OF TIMOTHY (PHLEUM PRATENSE)

Individual tillers of timothy ( Phleum pratense ) growing from seed in pots were labelled and their life history studied over a period of two years.
The leaf axil position of the first-produced tiller depended on its time of origin. A minimum of five leaves on the parent axis was necessary before its appearance, and no tiller emerged from the axil of any of the three terminal leaves of a shoot.
Ear-bearing capacity was high in early-formed tillers, but decreased progressively with later dates of origin despite favourable day length; all tillers arising after the end of July failed to produce inflorescences. A greater proportion of tillers on the main stem formed ears than secondary tillers of the same age. Time of ear emergence varied only within a few days for all tillers arising before the end of April. The number of leaves formed up to ear emergence varied linearly with date of origin and tiller position, the lowest recorded number being six.
Dry weight and length of inflorescence as well as seed yield per tiller varied linearly with the date of origin of tillers. Primary tillers generally had longer and heavier inflorescences and produced more seed per stem than secondary tillers of the same age.
Tillers arising at all times were found to have a limited span of life, not normally exceeding just over a year. The mean age of ear-bearing tillers varied linearly with their date of origin. Death-rates of all tillers generally increased with their age. Relative death-rates of fertile and vegetative tillers rose rapidly in the autumn but decreased during winter and early spring. Vegetative tillers which elongated in the year of formation survived for a shorter time than equivalent tillers in which stem elongation was delayed until the following year.
The perennial habit of grasses is discussed in relation to tiller survival. Successive new formation and death of tillers keep the plant in a condition of continuous dynamic change.     

Comparison of early development of three grasses: Lolium perenne, Agrostis stolonifera and Poa pratensis

BACKGROUND AND AIMS: To improve the management of grass communities, early plant development was compared in three species with contrasting growth forms, a caespitose (Lolium perenne), a rhizomatous (Poa pratensis) and a caespitose-stoloniferous species (Agrostis stolonifera). METHODS: Isolated seedlings were grown in a glasshouse without trophic constraints for 37 d (761 degrees Cd). The appearance of leaves and their location on tillers were recorded. Leaf appearance rate (LAR) on the tillers and site-filling were calculated. Tillering was modelled based on the assumption that tiller number increases with the number of leaves produced on the seedling main stem. Above- and below-ground parts were harvested to compare biomass. KEY RESULTS: Lolium perenne and A. stolonifera expressed similar bunch-type developments. However, root biomass was approx. 30 % lower in A. stolonifera than in L. perenne. Poa pratensis was rhizomatous. Nevertheless, the ratio of above-ground : below-ground biomass of P. pratensis was similar to that of L. perenne. LAR was approximately equal to 0.30 leaf d(-1) in L. perenne, and on the main stem and first primary tillers of A. stolonifera. LAR on the other tillers of A. stolonifera was 30 % higher than on L. perenne. For P. pratensis, LAR was 30 % lower than on L. perenne, but the interval between the appearance of two successive shoots from rhizomes was 30 % higher than the interval between two successive leaf stages on the main stem. Above-ground parts of P. pratensis first grew slower than in the other species to the benefit of the rhizomes, whose development enhanced tiller production. CONCLUSIONS: Lolium perenne had the fastest tiller production at the earliest stages of seedling development. Agrostis stolonifera and P. pratensis compensated almost completely for the delay due to higher LAR on tillers or ramets compared with L. perenne. This study provides a basis for modelling plant development.     

淹涝胁迫对水稻生育后期的生理特性和产量性状的影响

比较研究了水稻（湘中灿３号）在生育后期（孕穗期和乳熟期）进行没顶淹水胁拓处理后几个相关的生理指标和生长发育、产量的形态指标。研究结果显示水稻受淹后完全伸展叶片净光合速率降低,叶绿素含量在孕穗期明显降低,但在乳熟期只有少量下降。电导率明显增加,细胞膜透性增大,脯氨酸含量增加。奶系伤流量和,根系活力降低经与中群伤流液中的主要基酸如Ａｌａ,ＯＰｒｏ,Ｐｈｅ的含量增加,根系伤流量减少,根系活力降低,伤流液     

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.     

Life history and population dynamics of the Japanese Yam,Dioscorea japonica thunb.

Shoot morphology at the emergence ofDioscorea japonica Thumb. could be classed into the following three types: (1) a seedling emergence with only one leaf (Se type), (2) a plant consisting of one stem and one leaf, which has emerged from a small tuber (rhizophore) or bulbil less than 50 mg in dry weight (S type) and (3) a twiner with many leaves, which has emerged from a tuber or a bulbil of more than 50 mg in dry weight (L type). The Se type failed to develop beyond the second leaf stage in 1.5% sunlight exposure. The effects of initial plant (seeds, bulbils and tubers) size and light intensity on plant growth were analyzed. The larger the initial plant size was, the greater the growth in height and leaf area was. The distribution ratio of assimilated substances in leaves was high in smaller plants at the early growth stage. The distribution ratio in the tubers of larger plants became high at the early stage of growth. In all three types at over 3% sunlight exposure, the switch-over from the vegetative to reproductive growth phase occurred simultaneously at a later growth stage, but the Se type at 1.5% sunlight exposure showed a very early switch-over in its development; this switch-over may be related to shade tolerance capacity. The L type showed shade avoidance by forming a large productive structure as a twiner     

Developmental ideotype in the context of the dual‐purpose use of triticale,barley and durum wheat

Phenology strongly affects the performance of dual‐purpose (forage plus grain) cereals by defining the rate of leaf and tiller appearance, the duration of the pre‐grazing period and anthesis date. Here, the combined effect of genotypic and environmental variation conditions on anthesis date and the pre‐grazing period in durum wheat, triticale and barley was analysed to define a developmental ideotype for their dual‐purpose use. The extensive variation for the Haun stage reached at the time of apex emergence from the soil surface was associated with variation in the final leaf number, regardless of environment, species and cultivar. The timing of the terminal spikelet stage was the best predictor of apex emergence. The durum wheats tillered poorly, whereas the barleys and the triticales performed comparably in three of the four environments. In general, the higher the final leaf number, the higher the number of leaves and tillers produced by the time of apex emergence, the higher the number of leaves which emerges subsequently. The development of a high number of leaves both before and after apex emergence is advantageous, provided that anthesis is not overly delayed such that flowering and grain fill occur within a period where stressful conditions are likely to occur. Barley developed many leaves and had a low phyllochron, so was identified as the species with the development best suited for dual‐purpose cropping. However, the near simultaneous emergence of the first two tiller apices exposes it to a risk of grazing damage.     

Leaf Emergence of Spring Wheat Receiving Varying Nitrogen Supply at Different Stages of Development

We examined effects of nitrogen (N) supply on leaf emergenceof spring wheat (Triticum aestivum L.) grown in sand with nutrientsolution containing different N concentrations (9NO₃: 1NH₄).In expt 1, the cultivar 'Gamenya' received nutrient solutiontwice weekly containing a constant N supply ranging from 50to 2400 µM N. In expts 2 and 3, cultivars 'Aroona' and'Gamenya' were irrigated hourly with nutrient solution containingeither low (L = 50 µM N) or high (H = 2000 µM N)N supply. In expt 2, the N supply to half of the plants receivingL and H was changed at the double ridge stage of apical development,producing plants receiving LL, LH, HL and HH. In expt 3, N supplywas changed firstly when the main stem apex was vegetative (oneto two leaves) and secondly when the main stem apex was at doubleridge stage (four to five leaves), producing plants receivingLLL, LHL, HLH and HHH. Leaves on the main stem and primary tillerswere counted. Rate of leaf emergence was estimated from regressionof number of leaves against thermal time; the phyllochron wascalculated as 1/ rate of emergence. Severely N-deficient plants (which had at least a 60% reductionin shoot dry weight) had slower rates of leaf emergence (expt1). Fluctuating N supply sometimes, but not always, changedthe rate of leaf emergence (expts 2 and 3). The N supply beforedouble ridge stage had bigger effects on the phyllochron thanthat afterwards (expt 3). The phyllocrons of the main stemswere generally lower than those of tillers, with a greater differencebetween stems in N-deficient plants. Low N supply at the vegetativeapex stage decreased the total number of leaves on the mainstem, while low N supply after double ridge did not.Copyright1994, 1999 Academic Press Nitrogen, stress, spring wheat, Triticum aestivum, leaf emergence, phyllochron, apical development     

Leaf Blade Dimensions of Rice (Oryza sativa L. and Oryza glaberrima Steud.). Relationships between Tillers and the Main Stem

The effects of genotype and environment on the leaf area indexof rice are well documented, but the rules governing leaf areadistribution among main stem and tillers are less well understood.This study investigated the ontogenetic patterns of leaf bladearea and dimensions on the main stem and tillers of three ricecultivars, IAC47, Javaé (Oryza sativa L.) and CG14 (O.glaberrima Steud.) grown in pots without competition among plants.No differences in phyllochron were observed among culms, butleaves on tillers appeared 0.5 to 0.8 phyllochrons earlier thanthe genealogically corresponding leaves on the main stem (systemof Katayama). Cohorts were thus not fully synchronized. Allculms produced their largest leaves soon after panicle initiation(PI), despite differences in tiller age. Leaves that appearedsubsequently were smaller owing to reduced length but not width,which remained constant. Three different hypothetical rulesgoverning the behaviour of culms were tested: (1) tillers behaveas clones of the main stem; (2) cohorts behave uniformly, and(3) behaviour is determined by leaf position (sum of genealogicalleaf and tiller indices). The results were best explained byhypotheses (2) and (3), with leaf blade width and length showingmarkedly different patterns. Copyright 2001 Annals of BotanyCompany Phenology, phyllochron, leaf area, leaf position, leaf cohorts, summed leaf position, leaf blade length and width     

Distribution of imidacloprid residues in different parts of rice plants and its effect on larvae and adult females of Chilo suppressalis (Lepidoptera: Pyralidae)

The distribution of imidacloprid residue in different parts of rice, Oryza sativa L. (Graminales: Poaceae), plants was investigated for three rice varieties. Changes in larval hormones in Chilo suppressalis (Walker) (Lepidoptera: Pyralidae) that fed on imidacloprid-treated plant sources and the fecundity of resultant adult females (moths) also were determined. No significant differences in imidacloprid residue were detected in rice stem, leaves, and the unexpanded new leaf 7 d after treatment except in the rice variety Fengyouxiangzhan, in which residue content in leaves and the unexpanded new leaf was significantly lower than that in the stem. The percentage of reduction of imidacloprid residue after treatment varied with rice variety and plant parts. The greatest reduction in the three plant parts was shown in Wujing 15 compared with the other two varieties. The decomposition rate of imidacloprid residue in Wujing 15 was greater in leaves and unexpanded new leaf than in the stem, whereas in Wuyunjing 7 it was lower in leaves than in the unexpanded new leaf and stem. In Fengyouxiangzhan, the decomposition rate of imidacloprid in the stem increased 14 d after treatment. Juvenile hormone III and molting hormone titers in fourth instar larvae that had fed on rice plants treated with imidacloprid since the neonate stage were significantly higher than those in larvae fed on untreated plants (control). Juvenile hormone titer in larvae that had fed on Zizamia lalifolia Turcz galls treated with imidacloprid and the fecundity of the female moths that developed from such larvae were also significantly higher than that on the control counterparts. These results are discussed in relation to the impact of imidacloprid on nontarget insects and pest management.     

Phenotype/genotype associations for yield and salt tolerance in a barley mapping population segregating for two dwarfing genes

Barley traits related to salt tolerance are mapped in a population segregating for a dwarfing gene associated with salt tolerance. Twelve quantitative trait loci (QTLs) were detected for seven seedling traits in doubled haploids from the spring barley cross Derkado x B83-12/21/5 when given saline treatment in hydroponics. The location of QTLs for seedling growth stage (leaf appearance rate), stem weight prior to elongation, and tiller number are reported for the first time. In addition, four QTLs were found for the mature plant traits grain nitrogen and plot yield. In total, seven QTLs are co-located with the dwarfing genes sdw1, on chromosome 3H, and ari-e.GP, on chromosome 5H, including seedling leaf response (SGa) to gibberellic acid (GA(3)). QTLs controlling the growth of leaves (GS2) on chromosomes 2H and 3H and emergence of tillers (TN2) and grain yield were independent of the dwarfing genes. Field trials were grown in eastern Scotland and England to estimate yield and grain composition. A genetic map was used to compare the positions of QTLs for seedling traits with the location of QTLs for the mature plant traits. The results are discussed in relation to the study of barley physiology and the location of genes for dwarf habit and responses to GA.     

Isolation and characterization of a rice dwarf mutant with a defect in brassinosteroid biosynthesis

We have isolated a new recessive dwarf mutant of rice (Oryza sativa L. cv Nipponbare). Under normal growth conditions, the mutant has very short leaf sheaths; has short, curled, and frizzled leaf blades; has few tillers; and is sterile. Longitudinal sections of the leaf sheaths revealed that the cell length along the longitudinal axis is reduced, which explains the short leaf sheaths. Transverse sections of the leaf blades revealed enlargement of the motor cells along the dorsal-ventral axis, which explains the curled and frizzled leaf blades. In addition, the number of crown roots was smaller and the growth of branch roots was weaker than those in the wild-type plant. Because exogenously supplied brassinolide considerably restored the normal phenotypes, we designated the mutant brassinosteroid-dependent 1 (brd1). Further, under darkness, brd1 showed constitutive photomorphogenesis. Quantitative analyses of endogenous sterols and brassinosteroids (BRs) indicated that BR-6-oxidase, a BR biosynthesis enzyme, would be defective. In fact, a 0.2-kb deletion was detected in the genomic region of OsBR6ox (a rice BR-6-oxidase gene) in the brd1 mutant. These results indicate that BRs are involved in many morphological and physiological processes in rice, including the elongation and unrolling of leaves, development of tillers, skotomorphogenesis, root differentiation, and reproductive growth, and that the defect of BR-6-oxidase caused the brd1 phenotype.     

Rate of differentiation and emergence of nodal maize roots

The timing of root production is one of the parameters required for modelling the root system architecture. The objectives of this study are (1) to describe the rate of appearance of adventitious root primordia of maize and their rate of emergence out of the stem; (2) to test equations for the prediction of the rank of the phytomer on which root emergence occurs, in a wide range of field situations.Maize, cultivar Dea, was grown in controlled conditions and in the field in 1987, 1988, 1989 and 1991. Plants were regularly sampled and the following data were recorded: foliar stage, number of root primordia and number of emerged roots per phytomer. Root primordia were counted in transverse thin sections in the stem.At a single plant level, root primordia differentiation occurred sequentially on the successive phytomers, with no overlapping between two phytomers. The same was true for root emergence. Roots belonging to the same phytomer emerged at approximately the same time.At a plant population level, there was a linear relationship between the rank of the phytomer on which root primordia were differentiated and cumulated degree-days after sowing. A linear relationship was also observed between the rank of the phytomer on which roots were emerging and cumulated degree-days or foliar stage. In the range of field situations tested (several years, sowing dates and planting densities), both equations gave an accurate prediction of the timing of root emergence during the plant cycle.     

Photosynthesis and transpiration of the flag leaf in four spring-wheat cultivars

Co₂ exchange and transpiration rates of the flag leaves of four spring wheat (Triticum aestivum L.) cultivars, namely Glenlea, Neepawa, Opal and Kolibri, were compared using infra-red gas-analysis technique. The plants were grown in a controlled environment under an 18-h photoperiod, with day and night temperatures of 20 and 15° C, respectively. The time course of the CO₂-exchange rate (CER) of the flag leaf differed among cultivars. CER began to decrease rapidly some 2 weeks after ear emergence in Glenlea, Neepawa and Kolibri, but only after 4 weeks in Opal. The decline in CER of Glenlea, Neepawa and Opal was continuous throughout the period of grain development whereas in Kolibri CER was maintained at a constant level between the 4th and 6th weeks after ear emergence. The transpiration rates of the flag leaves of the 4 cultivars did not change markedly until 6–7 weeks after ear emergence, indicating that the reduction in CER was not primarily a response to increased stomatal resistance to the diffusion of CO₂. Removing the ear of the main shoot of intact plants failed to depress CER of the subtending flag leaf until 5 weeks after ear removal. Removing the ears of all the tillers of plants in which all but 3 tillers had been removed at ear emergence did not depress CER until 4 weeks after ear emergence, but removal of the ear of the main shoot of plants where all the tillers had been removed at ear emergence reduced the CER of the flag leaf 2 weeks after ear removal. Removal of tillers at ear emergence had a marked effect on the time course of CER and transpiration rates of the flag leaf. Both CER and transpiration rates of a 4-tiller plant were maintained at a higher level throughout ear development as compared to those of a one-tiller plant. The transpiration rate of the flag leaf of Glenlea increased during the later part of the life of the leaf even for one-tiller plants with no ear, indicating that such a stomatal response may be part of the normal course of leaf aging and not a response to a feedback stimulus from the ear.     

小麦不同蘖位叶片出生的两段线性模式及其品种、播期效应

叶片出生动态是小麦生长发育进程及其协调状况的重要表现,研究发现,小科叶片出生与播后累积ＧＤＤ（ｆｇｒｏｗｉｎｇ ｄｅｇｒｅｅ ｄａｙｓ ａｆｔｅｒ ｓｏｗｉｎｇ）的关系遵循两段（阶段Ⅰ快于阶段Ⅱ）线性模式,护颖分化期为两段模式的分界点,这一规律在正常发育的冬性和春性品种的７主茎及分蘖中表现一致,冬性品种播期１（９月３０日）、播期３（３月２日）的主茎及冬、春性品种各播期的Ｔ３分蘖,因生长发育异常而”     

Distribution patterns of assimilated 14C in vegetative and reproductive shoots of Lolium perenne and L. temulentum

The movement of ¹⁴C-labelled assimilate to the terminal meristem, stem, mature leaves, tillers and roots was measured in Loliurn perenn and Lolium temulentum after exposure to ¹⁴C0₂ of the youngest fully-expanded leaf and, on fewer occasions, the oldest healthy leaf on the main shoot. During early vegetative growth, the terminal meristem, tillers and roots received most of the ¹⁴C exported from the youngest leaf. As the shoot aged, more ¹⁴C was exported to the terminal meristem and tillers and less to roots. When the stem became a sizeable sink for ¹⁴C at the six-leaf (L. temulentum) or eleven-leaf (L. perenne) stage, less ¹⁴C moved to tillers and much less to roots. The terminal meristem continued to receive ¹⁴ at a steady rate throughout late vegetative growth. The transition from vegetative to reproductive growth in both species was marked by an abrupt increase in the export of ¹⁴C to stem from the upper leaf, but there was little change in the proportion of ¹⁴C which moved to the developing leaves and incipient inflorescence at the terminal meristem. At the same time, less ¹⁴C moved to tillers and much less to roots. Immediately before ear emergence, the export of ¹⁴C from the upper leaf (flag leaf) to the stem declined and the proportion moving to the ear increased, reaching a maximum of 55–75% as the ear emerged. The relative patterns of export of upper and lower leaves showed that while some ¹⁴ moved from each leaf to all meristems, the proximity of actively growing meristems appeared to be the main factor which determined the destination of most exported ¹⁴C. The distribution of ¹⁴C from upper and lower leaves was most alike in young vegetative plants of L. perenne. At later stages of development of both species, the terminal meristem and stem received most 14¹⁴C from the upper leaf, while roots and tillers received mos 14¹⁴C from the oldest leaf at the base of the shoot.     

Effect of nitrogen deficiency on photosynthesis and the partitioning of 14C-labelled leaf assimilate in unshaded and partially shaded plants of Lolium temulentum

The effect of a deficiency of applied nitrogen on the rate of leaf photosynthesis, and on the subsequent partitioning of ¹⁴C-labelled leaf assimilate between new leaf, stem, tillers and root, was investigated in single plants of Lolium temulentum L., grown normally in controlled environments, or grown with collars shading the base of the plant. The nitrogen deficiency reduced the rate of leaf photosynthesis, increased the retention of assimilate in the leaf, suppressed the export of assimilate to tillers, and generally increased the export of assimilate to roots and to new leaves. Shading the base of the plant generally had little effect on the rate of leaf photosynthesis, reduced the export of assimilate to roots, and increased the export of assimilate to new leaf and to the stem, which elongated when shading was imposed.