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.     

Supply and partitioning of assimilates to roots of Medicago sativa L. and Lotus corniculatus L. under anoxia

Abstract A current explanation of the mechanism of flooding injury to roots suggests that oxygen deficiency depresses the supply of respirable carbohydrates sufficiently to inhibit fermentation. However, even though it has been shown that phloem transport of assimilate is sharply reduced to anaerobic roots, inhibition of assimilate metabolism has also been suggested to be an important factor. This study examines these hypotheses by relating assimilate supply and metabolic activity in anoxic roots of alfalfa (Medicago sativa L.), a flood-intolerant species, and birdsfoot trefoil (Lotus corniculatus L.), a flood-tolerant plant. Roots were made anoxic (severe O₂ deficiency) for 2, 4 or 6 d and shoots were labelled with ¹⁴CO₂. Assimilate transport to the roots and metabolism to structural components were significantly decreased in both species in response to anoxia. Trefoil exhibited significantly greater ¹⁴C incorporation into the residue fraction at 4 d anoxia than did alfalfa, and this was consistent with the greater flooding tolerance of trefoil. When assimilate supply to O₂-deficient roots was decreased by shoot shading, shoot fresh weight was reduced by both anoxia and light treatments. Root-soluble sugars were significantly decreased by shading but were greatly increased in response to anoxia. Root starch concentration also increased under anoxia. Root K⁺ concentration was reduced by anoxia only. The energy status (ATP/ADP) of roots was significantly decreased by shading; however, anoxia reduced the energy status only in unshaded plants. The data indicate that carbohydrate supply to anaerobic roots does not appear to be a limiting factor in the metabolic response of alfalfa roots. Alternatively, metabolism of assimilate in anoxic roots may be an important determinant of survival.     

Defoliation and Regrowth in the Graminaceous Plant: The Role of Current Assimilate

Infra-red gas analysis and a quantitative radiocarbon tracertechnique were used to measure photosynthesis, and the export,distribution and utilization of current assimilate in the regrowthof leaf tissue and the growth of stem and root of partially-defoliateduniculm barley plants. After defoliation, which removed allleaf tissue above the ligule of leaf 3, the rate of photosynthesisof the remaining two older leaves fell to 90–95 per centof that of control leaves, but they exported more of their assimilatedcarbon to meristems elsewhere in the plant during the first48 h after the defoliation. The level of export from the twoolder leaves began to decline when new leaf tissue regrew fromthe shoot apex, and fell below that of the control leaves 4days after defoliation. The two older leaves supplied the assimilateused in the regrowth of new leaf tissue immediately after defoliation:previously they had exported most of their assimilate to root.There was no evidence that ‘reserves’ were mobilizedto meet the needs of regrowth at leaf meristems or, indeed,of the growth in stem and root; current photosynthesis suppliedsufficient assimilate to account for all observed growth. Ingeneral, the plants responded to defoliation with a rapid andmarked re-allocation of assimilate from root to leaf meristems,with the result that root growth was severely retarded but newleaf tissue grew at 70–100 per cent of the rate observedin control plants.     

半湿润地区氮磷钾配施对强筋小麦功能叶光合速率的影响

选用优质强筋小麦品种陕253为材料,研究了不同氮磷钾肥水平及组合对小麦生育后期主茎、主茎分蘖Ⅰ和主茎分蘖Ⅱ功能叶净光合速率的影响,结果表明:在不同的NPK肥配比处理中,功能叶净光合速率在蘖位、叶位间差异均极其显著,不同生育期间存在明显差异,具体表现为:孕穗期高于灌浆期,主茎>主茎分蘖Ⅰ>主茎分蘖Ⅱ,旗叶高于倒二叶.孕穗期以N225P120K120处理对茎蘖功能叶光合速率影响最为明显,分别比CK1和CK2增加20.41%和25.76%;灌浆期以N135P225K120处理对茎蘖旗叶净光合速率作用显著,分别比CK1和CK2增加43.48%和10.83%.研究发现,在一定磷钾肥基础上通过调节氮肥可以调节孕穗期茎蘖功能叶净叶光合速率,提高钾肥用量有利于提高灌浆期各茎蘖功能叶的净光合速率.     

Assimilate Distribution in Poa annua L.

The carbon economy of a flowering tiller of Poa annua L. hasbeen examined over the period from inflorescence emergence tograin shedding. The total import of ¹⁴C by the inflorescencereached a maximum at late grain filling but the relative importof assimilate was greatest 14 days after its appearance andrepresented 20–25 per cent of that assimilated by theinflorescence itself. The inflorescence continued to be an importantassimilatory organ after grain ripening when it exported morethan 50 per cent of its assimilate to the stem, roots and othertillers. The patterns of distribution of assimilates from the youngestuppermost and the oldest green leaf of the reproductive tillerwere largely determined by the stage of development of the inflorescence.The youngest leaf mainly supported the inflorescence up to theend of the grain-filling stage but then supplied assimilatesbasally to the roots and adjacent tillers. The oldest greenleaf supported the growth of the stem and the inflorescenceup to anthesis but after this supplied assimilates mainly tothe roots and tillers. Removal of grains or the entire inflorescence only 1 h beforesupplying ¹⁴CO₂ greatly reduced the rate of fixation of ¹⁴CO₂and the export of radiocarbon, as well as changing the patternof distribution of assimilates within the plant. The significanceof these results is discussed and comparisons made with cerealsand perennial grasses.     

Interactive effects of elevated atmospheric CO2, mycorrhization and drought on long-distance transport of reduced sulphur in young pedunculate oak trees (Quercus robur L.)

Pedunculate oak (Quercus robur L.) was germinated and grown under nutrient non-limiting conditions for a total of 10–15 weeks at ambient CO₂ concentration and 1100 μmol mol^–1 CO₂ either in the presence or the absence of the mycorrhizal fungus Laccaria laccata. Half of the oak trees of these treatments were exposed to drought during final growth by suspending the water supply for 21 d. Mycorrhization and elevated atmospheric CO₂ each enhanced total plant biomass per tree. Whereas additional biomass accumulation of trees grown under elevated CO₂ was mainly attributed to increased growth of lateral roots, mycorrhization promoted shoot growth. Water deficiency reduced biomass accumulation without affecting relative water content, but this effect was more pronounced in mycorrhizal as compared to non-mycorrhizal trees. Elevated CO₂ partially prevented the development of drought stress, as indicated by leaf water potential, but did not counteract the negative effects of water deficiency on growth during the time studied. Enhanced biomass accumulation requires adaption in protein synthesis and, as a consequence, enhanced allocation of reduced sulphur produced in the leaves to growing tissues. Therefore, allocation of reduced sulphur from oak leaves was studied by flap-feeding radiolabelled GSH, the main long-distance transport form of reduced sulphur, to mature oak leaves. Export of radiolabel proceeded almost exclusively in basipetal direction to the roots. The rate of export of radioactivity out of the fed leaves was significantly enhanced under elevated CO₂, irrespective of mycorrhization. A higher proportion of the exported GSH was transported to the roots than to basipetal stem sections under elevated CO₂ as compared to ambient CO₂. Mycorrhization did not affect ³⁵S export out of the fed leaves, but the distribution of radiolabel between stem and roots was altered in preference of the stem. Trees exposed to drought did not show appreciable export of the ³⁵S radioactivity fed to the leaves when grown under ambient CO₂. Apparently, drought inhibited basipetal transport of reduced sulphur at the level of phloem loading and/or phloem transport. Elevated CO₂ seemed to counteract this effect of drought stress to some extent, since higher leaf water potentials and improved ³⁵S export out of the fed leaves was observed in oak trees exposed to drought and elevated CO₂ as compared to trees exposed to drought and ambient CO₂.     

Carbon dioxide exchange in relation to sink demand in wheat

Summary In this paper, experiments are described which examine the effect of requirement for assimilates by the ear on the rate of net photosynthesis in leaves of wheat (Triticum aestivum L.). Different levels of requirement were achieved by various levels of sterilization of florets just before anthesis, which resulted in a range of grain numbers per ear, and by inhibiting photosynthesis of the intact ear by 3-(3,4-dichlorophenyl)-1,1-dimethylurea (DCMU). Only the ear and two uppermost leaves of the main shoot were considered, all the lower leaves and tiller leaves being excised when the experimental treatments were imposed. In two experiments, tiller regrowth was permitted during the experimental period, while in a third, new tillers were defoliated regularly.The response of leaf photosynthesis to the level of assimilate requirement by the ear was influenced by the treatment of the vegetative tillers. Thus, the net photosynthesis rate of the flag leaf was decreased by a reduction in grain number, or increased by inhibition of photosynthesis in the ear, only when the vegetative tillers were kept defoliated; when these tillers were allowed to re-grow normally, there was no influence of ear treatment on leaf photosynthesis. Temporal changes in leaf photosynthesis were consistent with this response pattern, i.e., when tillers were defoliated, the initial high rates of photosynthesis persisted for much longer.In the experiment where photosynthesis was influenced by the requirement for assimilate in the ear, the variation occurred through change in stomatal conductance on the abaxial surface of the leaf. This surface has a lesser conductance to CO₂ exchange than the adaxial surface. The implication of this finding to rapid methods of plant screening is discussed.     

Analysis of assimilate distribution by 3H-tracing during recovery period after rice seedling transplanting

In order to characterize assimilate distribution in rice (Oryza sativa L.) seedlings during recovery period after transplanting, the experiments were conducted by using ³H-isotope tracer technique with three rice hybrid combinations. The label introduced into the leaf sheath was distributed and redistributed to young roots, tillers, and young leaves, thus promoting these organs to grow at the initial stage after transplanting. During period from 10 to 30 days after labeling, ³H-assimilate redistribution occurred mainly from the sheath of the main stem; some radioactivity moved to tillers and roots emerged after transplanting. The activity and percentage of ³H-compounds transporting to rooting nodes and roots emerged after transplanting were higher in root-pruned plants than in control, indicating that growth of root-pruned seedlings more depended on the export of the compounds from the sheath of the main stem than growth of control seedlings.     

The Effect of Nitrogen Supply to Urtica dioica L. Plants on the Distribution of Assimilate Between Shoot and Roots

In this study the influence of nitrogen nutrition on the patterns of carbon distribution was investigated with Urtica dioica. The nettles were grown in sand culture at 3 levels of NO^?₃, namely 3 (low), 15 (medium) and 22 (high) mM. These levels encompassed a range within which nitrogen did not affect total biomass production. The ratio of root: shoot biomass of the low nitrogen plants was, however, significantly higher than that of the nettles grown at medium and high N supply. Carbon allocation from one leaf of each pair of leaves was examined after a ¹⁴CO₂-pulse and a subsequent ¹⁴C distribution period of one night. Only the youngest two leaf pairs did not export assimilates. Carbon (¹⁴C) export to the shoot apex and to the roots, as measured at the individual nodes responded to the nitrogen status: At medium and high nitrogen supply the 3rd, 4th and 5th leaf pairs exported to the shoot apex, while lower leaves exported to the root. At low nitrogen supply only the 3rd leaf exported towards the shoot apex. The results illustrate the plastic response of carbon distribution patterns to the nitrogen supply, even when net photosynthesis, carbon export from the source leaves and biomass production were not affected by the nitrogen supply to the plant.     

A Quantitative Analysis of the Uptake of Carbon and of the Supply of 14C-labelled Assimilates to Areas of Meristematic Growth in Lolium temulentum

A quantitative analysis of the ¹⁴C-labelled assimilate suppliedby leaves on the main shoot to terminal meristem, stem, tillers,and roots was conducted during parallel periods of reproductiveand vegetative development in Lolium temulentum. The initial rate of entry of carbon into the shoot varied withthe area and photosynthetic efficiency of the assimilating leaf.Subsequently, respiratory losses of carbon during translocationand incorporation of assimilate at the site of utilization alsovaried. The combined effect of these differences resulted inthe supply of recently assimilated carbon being twofold greaterin reproductive shoots than in vegetative shoots, while withinshoots the carbon supply of the youngest fully expanded leafranged from four-or five-fold greater than the oldest leaf inyoung shoots, to two-or three-fold greater in older shoots.In both reproductive and vegetative shoots, the two or threeyoungest leaves thus dominated the supply of carbon for meristematicgrowth. Meristematic tissue in expanding leaves and leaf primordia atthe terminal meristem of the vegetative shoot received 18–27per cent of the total shoot carbon. This meristem utilized aboutthe same proportion of shoot carbon when it developed into aninflorescence, indicating no major change in the level of meristematicactivity. The proportion of shoot carbon utilized in stem growthincreased as both reproductive and vegetative shoots aged; thisincreased meristematic activity in stem internodes was accompaniedby reduced export of carbon to roots, which received less than10 per cent of the shoot carbon when the experiments ended.The main shoot translocated 20–30 per cent of its recentlyassimilated carbon to developing and rooted tillers, which assinks for carbon were thus as important as the terminal meristemand stem. This outward flow of carbon continued relatively uncheckedwhen donor and receptor shoots developed inflorescences.     

Effects of geminivirus infection and growth irradiance on the vegetative growth and photosynthetic production of Eupatorium makinoi

Eupatorium makinoi plants with or without geminivirus infection were grown in shading frames with 70, 15 and 5.5% sunlight. Growth characteristics of these plants in the early vegetative phase were compared by means of growth analysis. We also measured leaf photosynthetic gas exchange rates and examined relationships between leaf photosynthesis and whole-plant growth. Relative growth rate (RGR=(1/W)×(dW/dt), where W is plant dry mass) of virus-infected plants was lower than that of uninfected plants under all three light conditions. The reduction of RGR by infection was increased with irradiance. The net assimilation rate (NAR=(1/A)×(dW/dt), where A is total leaf area of the plant) was also reduced both by infection and shading. NARs that were estimated from light-response curves of leaf photosynthesis, in situ measurements of irradiance, and respiration rates of leaves, stems and below-ground parts, agreed very well with the values obtained by conventional growth analysis techniques. Decreases in the estimated NAR value from infection and shading were mostly explained by the decreases in leaf photosynthesis. These results clearly showed that lowered RGR in virus-infected plants was attributed mainly to impaired photosynthesis in virus-infected leaves.     

The photosynthesis of ryegrass leaves grown in a simulated sward

Plants were taken from simulated swards of perennial ryegrass (Lolium perenne) grown in a controlled environment and the rates of photosynthesis of the youngest fully expanded leaves, and the second and third youngest leaves on the same tillers were measured. The youngest leaves had the highest rates and the third the lowest, with the second leaves intermediate. The rate of photosynthesis in bright light of successive youngest expanded leaves decreased as the swards increased in leaf area, but did not when plants were grown so that the main stem was not shaded. When plants were grown at different densities and the photosynthetic rates of leaves of a particular ontogenetic rank were measured, it was found that leaves on plants from higher densities had lower rates of photosynthesis. Also leaves on plants grown in bright light had higher photosynthetic rates than those on plants grown in dim light. It is concluded that the decline in the photosynthetic capacity of successive leaves in a rapidly growing simulated sward is due to the intense shading to which they are subjected during their expansion.     

Nitrogen reallocation and photosynthetic acclimation in response to partial shading in soybean plants

The first trifoliate of soybean was shaded when fully expanded, while the plant remained in high light; a situation representative for plants growing in a closed crop. Leaf mass and respiration rate per unit area declined sharply in the first few days upon shading and remained rather constant during the further 12 days of the shading treatment. Leaf nitrogen per unit area decreased gradually until the leaves were shed. Leaf senescence was enhanced by the shading treatment in contrast to control plants growing in low light. Shaded leaves on plants grown at low nutrient availability senesced earlier than shaded leaves on plants grown at high nutrient availability. The light saturated rate of photosynthesis decreased also gradually during the shading treatment, but somewhat faster than leaf N, whereas chlorophyll contents declined somewhat slower than leaf N.
Partitioning of N in the leaf over main photosynthetic functions was estimated from parameters derived from the response of photosynthesis to CO₂. It appeared that the N exported from the leaf was more at the expense of compounds that make up photosynthetic capacity than of those involved in photon absorption, resulting in a change in partitioning of N within the photosynthetic apparatus. Photosynthetic nitrogen use efficiency increased during the shading treatment, which was for the largest part due to the decrease in leaf N content, to some extent to the decrease in respiration rate and only for a small part to change in partitioning of N within the photosynthetic apparatus.     

An appraisal of ear, leaf and stem 14CO2 assimilation, 14C-assimilate distribution and growth in a reproductive seed crop of amenity Lolium perenne

In seed crops of Lolium perenne, yield may be reduced by competition for a limited assimilate supply from sinks other than the ear. This study was undertaken to evaluate the priorities for allocation of assimilate within the crop from all photosynthetic sites on the main reproductive tiller after anthesis. Ear, stem and leaves were fed with ¹⁴CO₂ on two occasions; the assimilatory efficiency of these sources and the magnitude and pattern of ¹⁴C-assimilate export from each was determined. The growth of each part of the main tiller and subtending tillers was also measured. Stem elongation apparently dominated the current assimilate resource and the ear did not become a net importer of assimilate until this process had ceased. Assimilate allocation to the tillers was high throughout. The nature of any competition between these sinks is discussed. When crop growth was contrasted with that in a previous year, environmental factors were implicated as determinants of priority for assimilate allocation to each sink. Sources of carbon for seed filling are also discussed as is the relevance of these findings to seed crop management.     

Plant leaf area expansion and assimilate production in wheat (Triticum aestivum L.) growing under low phosphorus conditions

Under phosphorus deficiency reductions in plant leaf area have been attributed to both direct effects of P on the individual leaf expansion rate and to a reduced availability of assimilates for leaf growth. In this work we use experimental and simulation techniques to identify and quantify these processes in wheat plants growing under P-deficient conditions. In a glasshouse experiment we studied the effects of soil P addition (0–138 kg P₂O₅ ha^-1) on tillering, leaf emergence, leaf expansion, plant growth, and leaf photosynthesis of wheat plants (cv. INTA Oasis) that were not water stressed. Plants were grown in pots containing a P-deficient (3 mg P g^-1 soil) sandy soil. Sowing and pots were arranged to simulate a crop stand of 173 plants m^-2. Experimental results were integrated in a simulation model to study the relative importance of each process in determining the plant leaf area during vegetative stages of wheat. Phosphorus deficiency significantly reduced plant leaf area and dry weight production. Under P-deficient conditions the phyllochron (PHY) was increased up to a 32%, compared to that of high-P plants. In low-P plants the rate of individual leaf area expansion during the quasi-linear phase of leaf expansion (LER) was significantly reduced. The effect of P deficiency on LER was the main determinant of the final size of the individual leaves. In recently expanded leaves phosphorus deficiency reduced the photosynthesis rate per unit leaf area at high radiation (AMAX), up to 57%. Relative values of AMAX showed an hyperbolic relationship with leaf P% saturating at 0.27%. Relative values of the tillering rate showed an hyperbolic relationship with the shoot P% saturating at values above 0.38%. The value of LER was not related to the concentration of P in leaves or shoots. A morphogenetic model of leaf area development and growth was developed to quantify the effect of assimilate supply at canopy level on total leaf area expansion, and to study the sensitivity of different model variables to changes in model parameters. Simulation results indicated that under mild P stress conditions up to 80% of the observed reduction in plant leaf area was due to the effects of P deficiency on leaf emergence and tillering. Under extreme P-deficient conditions the simulation model failed to explain the experimental results indicating that other factors not taken into account by the model, i.e. direct effects of P on leaf expansion, must have been active. Possible mechanisms of action of the direct effects of P on individual leaf expansion are discussed in this work.     

Photosynthesis and carbon export in white clover plants grown at various levels of phosphorus supply

Net photosynthesis, concurrent carbon export and starch, sucrose and inorganic phosphorus concentrations were measured in leaves of white clover ( Trifolium repens L. cv. Grasslands Huia) grown at four levels of phosphorus supply in the presence or absence of mineral nitrogen. The nitrogen treatments had no effect on growth, photosyntheis or carbon export. At the three higher levels of phosphorus supply, the amount of carbon exported was about 77% of net fixation. Photosynthesis and export per leaf decreased with phosphorus supply, primarily through the effect of phosphorus supply on leaf area. The rate of photosynthesis was reduced only at the lowest level of phosphorus supply.
Inorganic phosphorus rose with phosphorus supply but starch concentration was unaffected. Sucrose was reduced at the lowest level of phosphorus supply but not significantly affected at higher levels. The ratio between starch and sucrose concentration was also unaffected at the higher levels, but was increased at the lowest level of supply. There thus appeared to be direct effects of phosphorus supply on photosynthesis, partitioning of carbon to carbohydrates and, by implication, export, only at the lowest level of phosphorus supply. As leaf area and plant growth were affected over the whole range of phosphorus supply, factors other than photosynthesis per se must have determined the response of growth to phosphorus supply.     

Allocation responses to CO2 enrichment and defoliation by a native annual plant Heterotheca subaxillaris

Among plants grown under enriched atmospheric CO₂, root:shoot balance (RSB) theory predicts a proportionately greater allocation of assimilate to roots than among ambient‐grown plants. Conversely, defoliation, which decreases the plant's capacity to assimilate carbon, is predicted to increase allocation to shoot. We tested these RSB predictions, and whether responses to CO₂ enrichment were modified by defoliation, using Heterotheca subaxillaris, an annual plant native to south‐eastern USA. Plants were grown under near‐ambient (400 μmol mol^?1) and enriched (700 μmol mol^?1) levels of atmospheric CO₂. Defoliation consisted of the weekly removal of 25% of each new fully expanded, but not previously defoliated, leaf from either rosette or bolted plants. In addition to dry mass measurements of leaves, stems, and roots, Kjeldahl N, protein, starch and soluble sugars were analysed in these plant components to test the hypothesis that changes in C:N uptake ratio drive shifts in root:shoot ratio. Young, rapidly growing CO₂‐enriched plants conformed to the predictions of RSB, with higher root:shoot ratio than ambient‐grown plants (P < 0.02), whereas older, slower growing plants did not show a CO₂ effect on root:shoot ratio. Defoliation resulted in smaller plants, among which both root and shoot biomass were reduced, irrespective of CO₂ treatment (P < 0.03). However, H. subaxillaris plants were able to compensate for leaf area removal through flexible shoot allocation to more leaves vs. stem (P < 0.01). Increased carbon availability through CO₂ enrichment did not enhance the response to defoliation, apparently because of complete growth compensation for defoliation, even under ambient conditions. CO₂‐enriched plants had higher rates of photosynthesis (P < 0.0001), but this did not translate into increased final biomass accumulation. On the other hand, earlier and more abundant yield of flower biomass was an important consequence of growth under CO₂ enrichment.     

The Measurement and Prediction of Organ Growth in a Uniculm Barley

Measurements of leaf areas, net rates of photosynthesis, patternsof assimilate translocation, and of some aspects of respirationwere made at leaf-increment intervals during the expansion ofleaves 5, 6, 7, 8, and 9 on the single axis of a uniculm barley(Kindred Uniculm 97) grown in controlled environments. Thesedata were used as the primary inputs in a computer programmedeveloped to simulate the carbon metabolism and consequent weightchanges of the organs in the single-axis barley plant. The totalweight of plant tissue increased threefold between the expansionof the fifth and ninth leaves; during this period the simulationmodel generally predicted the daily growth increments to within10 per cent of the observed values. The predictions of dailygrowth increments in new leaf, stem, and root were less accurate.The simulation indicated that the proportion of photosyntheticproducts incorporated in new growth at the meristems declinedfrom some 54 per cent of the assimilate at the fifth leaf stageto 42–3 per cent at the ninth leaf stage. This declinein the efficiency of conversion of photosynthetic products appearedto be the result of an increase in maintenance respiration,which in turn stemmed from an approximately linear increasein total tissue weight; the proportion of photosynthetic productslost in the respiration associated with synthetic processesremained approximately constant throughout the growth period.     

The patterns of growth, assimilation of 14CO2 and distribution of 14C-assimilate within vegetative plants of Loliumperenne at low and high density

The patterns of growth, assimilation of ¹⁴CO₂ and distribution of ¹⁴C-labelled assimilate were followed for 12 wk from sowing in individual plants of Lolium perenne grown in miniswards at either low (500 plants m^-2) or high (5000 plants m^-2) density. At the latter density, plants were characterised by a 50% reduction in RGR, by the production of fewer tillers, especially second- and third-order tillers, and by a reduction in mean tiller weight. All the green and senescing leaves of each tiller assimilated ¹⁴CO₂ and the overall assimilatory capacity of a tiller was directly related to its dry weight. At both densities the plant consisted of a main shoot and established tillers with comparable assimilatory activities and a range of developing tillers that assimilated relatively small amounts of ¹⁴CO₂. As each successive primary tiller developed it was supplied with assimilate from the main shoot and the degree of support was inversely proportional to the dry weight of the tiller. At both densities it was concluded that the first primary tiller could be regarded as an independent assimilatory unit when it reached a dry weight of about 25 mg even though some import of main shoot assimilate continued until the tiller was double this weight. The supply of assimilate to the root system was greatly reduced at both densities compared with previous observations on plants grown singly.     

Carbon allocation to shoots and roots in relation to nitrogen supply is mediated by cytokinins and sucrose: Opinion

In this paper we firstly show some general responses of biomass partitioning upon nitrogen deprivation. Secondly, these responses are explained in terms of allocation of carbon and nitrogen, photosynthesis and respiration, using a simulation model. Thirdly, we present a hypothesis for the regulation of biomass partitioning to shoots and roots.Shortly after nitrogen deprivation, the relative growth rate (RGR) of the roots generally increases and thereafter decreases, whereas that of the shoot decreases immediately. The increased RGR of the root and decreased RGR of the shoot shortly after a reduction in the nitrogen supply, cause the root weight ratio (root weight per unit plant weight) to increase rapidly.We showed previously that allocation of carbon and nitrogen to shoots and roots can satisfactorily be described as a function of the internal organic plant nitrogen concentration. Using these functions in a simulation model, we analyzed why the relative growth rate of the roots increases shortly after a reduction in nitrogen supply. The model predicts that upon nitrogen deprivation, the plant nitrogen concentration and the rate of photosynthesis per unit plant weight rapidly decrease, and the allocation of recently assimilated carbon and nitrogen to roots rapidly increases. Simulations show that the increased relative growth rate of the root upon nitrogen deprivation is explained by decreased use of carbon for root respiration, due to decreased carbon costs for nitrogen uptake. The stimulation of the relative growth rate of the root is further amplified by the increased allocation of carbon and nitrogen to roots. Using the simple relation between the plant nitrogen concentration and allocation, the model describes plant responses quite realistically.Based on information in the literature and on our own experiments we hypothesize that allocation of carbon is mediated by sucrose and cytokinins. We propose that nitrogen deprivation leads to a reduced cytokinin production, a decreased rate of cytokinin export from the roots to the shoot, and decreased cytokinin concentrations. A reduced cytokinin concentration in the shoot represses cell division in leaves, whereas a low cytokinin concentration in roots neutralizes the inhibitory effect of cytokinins on cell division. A reduced rate of cell division in the leaves leads to a reduced unloading of sucrose from the phloem into the expanding cells. Consequently, the sucrose concentration in the phloem nearby the expanding cells increases, leading to an increase in turgor pressure in the phloem nearby the leaf's division zone. In the roots, cell division continues and no accumulation of sugars occurs in dividing cells, leading to only marginal changes in osmotic potential and turgor pressure in the phloem nearby the root's cell division zone. These changes in turgor pressure in the phloem of roots and sink leaves affect the turgor pressure gradients between source leaf-sink leaf and source leaf-root in such a way that relatively more carbohydrates are exported to the roots. As a consequence RWR increases after nitrogen deprivation. This hypothesis also explains the strong relationship between allocation and the plant nitrogen status.