The Distribution of Assimilates in Lolium multiflorum Lam. Following Differential Defoliation

In seedling plants of Lolium multiflorum Lam. the tillers weredefoliated but the main shoot was left intact. Radiocarbon as¹⁴CO₂ was supplied to this shoot at different times followingtiller defoliation and the pattern of distribution of labelledassimilates was determined quantitatively. It was found thata greater proportion (approximately 10–20 per cent) ofexported assimilate was translocated to the cut tillers butalthough the proportion supplied to the root system was lessthe total radiocarbon incorporated by the roots was unchanged.This was brought about by a large increase in the export ofradiocarbon fixed by the intact shoot—up to 100 per centfollowing one treatment. These alterations in the organizationof the defoliated plant lead to a greater efficiency in thecarbon economy and are discussed in relation to the stress imposedby defoliation.     

Sugar Accumulation in Barley and Rice Grown in Solutions with low Concentrations of Oxygen

Barley and rice, at the early tillering stage, were grown inaerated nutrient solutions (> 7 mg O₂ l^–1) and transferredto solutions of low O₂ concentrations (< 0.5 mg l ^–1). For barley, low O₂ concentrations during the first 5 days severelyinhibited growth of seminal roots had less effect on nodal roots,and did not reduce shoot growth. Longer exposure to low O₂ concentrationsreduced shoot as well as root growth. Sugar concentrations inroots and shoots increased within 7 h after transfer of plantsto low O₂ concentrations. After 5 days at low O₂ concentrationssugar concentrations were very high in fast growing nodal rootsand in shoots, as well as in the slower growing seminal roots. In rice, low O₂ concentrations increased sugar levels of rootsduring summer, but not during winter. In summer, the highersugar levels at low O₂ concentrations persisted throughout adiurnal cycle. In root apices, sugar concentrations were increasedby low O₂ concentrations, even though the experiment was donein winter and the bulk of the root system showed no differencein sugar levels. The data indicate that sugar accumulation, at low O₂ concentrations,is caused by reduced growth and also that even apices of rootsgrown at low O₂ concentrations have sufficient substrates forrespiration. Hordeum vulgare L, barley, Oryza sativa L, rice, sugar accumulation, oxygen concentration     

Root growth and nitrate uptake by wheat (Triticum aestivum L.) following wetting of dry surface soil

The nitrate uptake capacity of surface roots of spring wheat(Triticum aestivum L. cv. Kulin) was investigated followingwetting of dry surface soil. Plants were grown to stem elongationstage with adequate watering at depth while the surface soilwas allowed to dry. Eight weeks after sowing, water or a ¹⁵N-nitratesolution was added to the surface soil to simulate rainfall.Root growth and nitrate uptake were measured up to 4 d afterwetting on plants with unconfined nodal root growth and on plantswith the majority of nodal roots confined within small vials.Prior to wetting, plants from both nodal treatments had seminalroots with collapsed cortices along the upper 10 cm and manyshort, viable lateral roots. Nodal roots, where present, wereonly a few cm long and unbranched. Only plants with unconfinednodal roots were able to take up nitrate within the 24 h beforeany new root growth. By 2 d after wetting there was significantgrowth of the seminal lateral roots, and rapid growth and branchingof nodal roots. From 2 d after wetting, plants with confinednodal roots also took up nitrate, presumably due to the growthof the seminal lateral roots. Hence it appears as though thenodal roots in the unconfined treatment could immediately takeup nitrate, but the seminal roots required new lateral rootgrowth to become active in nitrate uptake. The plants with confinednodal roots had a lower nitrate uptake than those with unconfinednodal roots 4 d after wetting, indicating that the seminal rootsystem was not able to compensate for lack of nodal roots. Insufficientnitrate was taken up after 4 d, by plants from either nodalroot treatment, to increase the shoot N concentration significantly. Key words: Triticum aestivum, nitrate uptake, drought, seminal roots, nodal roots     

Patterns of 14C-labelled Assimilate Partitioning in Red and White Clover During Vegetative Growth

A quantitative analysis of the ¹⁴C-labelled assimilate suppliedby the expanded leaves on the primary shoot to growing leaves,stem, lateral shoots (branches or stolons) and roots in redand white clover was conducted during vegetative growth. Stem growth of the primary shoot was inhibited in both cloversand utilized no energy resources. The growing leaves at theprimary shoot apex of white clover imported 4 per cent of theshoot's assimilate compared with 10 per cent in red clover.At the basal end of the primary shoot, the tap root of whiteclover imported 16 per cent of the shoot's assimilate comparedwith 22 per cent in red clover. Branches in red clover and stolonsin white clover were by far the largest sinks for primary shootassimilate, importing 39 per cent and 63 per cent of the labelledassimilate, respectively. Analyses of the translocation of assimilate from individualprimary shoot leaves demonstrated that in both clovers olderleaves exported more of their assimilate to branches or stolons,whereas younger leaves exported more of their assimilate toroots, and possibly in white clover, to growing leaves at thetip of the shoot. Of the labelled assimilate exported to branchesor stolons, each primary shoot leaf exported preferentiallyto the branch or stolon in its own axil, but in addition exportedsubstantial quantities of assimilate to all other axillary shoots,particularly those arising from basal axils where the subtendingleaf had died. Trifolium repens, Trifolium pratense, red clover, white clover, assimilate partitioning, perennation     

Carbon Economy of Carrots During Initiation of the Storage Root in Cultivars Contrasting in Shoot: Root Ratio at Maturity

Photosynthesis, respiration and growth of two cultivars of carrotwith contrasting ratios of shoot: storage root weight at maturity,were compared during initiation of the storage root at 20 °C.Partition of assimilate between shoot, roots and respirationshowed no varietal differences but distribution between storageand fibrous roots was different from the time that the storageroot could be morphologically identified. For both cultivarsover the period investigated, approximately 64% of net photosynthesiswas partitioned to the shoot with 5% lost as respiration duringthe dark and 59% used in growth. Of that exported to the rootsystem (36%), 19% (of net photosynthesis) was used in growthand 17% was lost in respiration. In the cultivar with greatershoot: storage root ratio at maturity, 4.6% was allocated tothe storage root in contrast to 7.5% in the cultivar with alesser shoot: storage ratio at maturity. It is concluded thatgreater dry matter accumulation in the storage root of the lattercultivar does not result from transient differences in respiratoryloss and is not evident in shoot to total root dry matter distributionover this period. Daucus carota L, carrot, assimilate partition, shoot, storage root, shoot: root ratio     

The Reciprocal Transfer of Radiocarbon between a Developing Tiller and its Parent Shoot in Vegetative Plants of Lolium multiflorum Lam.

The transfer of ¹⁴C-labelled assimilates between a tiller andits parent shoot was examined in young plants of Lolium multiflorumLam. Radiocarbon was exported freely from an expanded laminato sinks within the shoot axis from which it originated andto the root system. Lesser amounts of radiocarbon were exportedto the other shoot. It is suggested that the reciprocal exchangeof radiocarbon between tiller and main shoot occurred principallyvia a direct pathway through stem tissues rather than via apathway involving the roots.     

The Interdependence of Tillers in Lolium multiflorum Lam.--a Quantitative Assessment

Experiments were made to determine the extent of reciprocaltransfer of products (derived from the assimilation of ¹⁴CO₂)between various parts of the young vegetative grass plant (Loliummultiflorum Lam.). When individual laminae on different tillerswere supplied with ¹⁴CO₂, 47–64 per cent of the fixedcarbon was exported after 24 h. The principal sinks were theroot system and the shoot or tiller to which the fed leaf wasattached. Other tillers also received significant quantitiesof radiocarbon. When whole tillers were supplied with ¹⁴CO₂the percentage of fixed radiocarbon exported within 24 h rangedfrom 14–31 per cent. Of this, 50–74 per cent wasrecovered from the root system (except in the case of exportfrom the youngest tiller) but exchange of material between tillersalso occurred.A reciprocity diagram is presented and it is concludedthat despite the magnitude of the exchange no tiller showedan over-all net gain or loss. The main shoot and the tillersdiffered in the extent of their carbon exchange and in theirdegree of independence. The oldest daughter tiller of the mainshoot was the most independent and the main shoot most interdependent.     

Partition of 14C Assimilate between Organs and Fractions of Contrasting Varieties of Carrot during Initiation of the Storage Root

This work examines the differences in partition and activityof ¹⁴C in two varieties of carrot (Daucus carota L.) contrastingin shoot to storage root ratio at maturity. Plants were grownin a controlled environment of 20 ?C and 500 µmol m^–2s^–1. During initiation of the storage root (10–25d from sowing) plants were exposed to ¹⁴CO₂ for 1 h and theradioactivity in ethanol-soluble and -insoluble fractions ofshoots, storage and fibrous roots estimated at various timesup to 48 h after exposure. Between 35% and 40% of radioactivityinitially present in the plants was respired during the first24 h and 25–35% of that remaining after 24 h was foundin the roots, depending on age. The proportion found in thestorage region remained fairly constant between 15 and 25 dand was smaller than at 10 d. In the variety with a larger proportionof storage root at maturity (cv. Super Sprite), there was agreater proportion of label in both ethanol-soluble and -insolublefractions of the storage region soon after storage root initiationhad begun than in the variety with a smaller proportion of storageroot at maturity (cv. Kingston). There was no varietal differencein specific activities of the storage roots, but fibrous rootsof cv. Super Sprite showed a greater specific activity thanin cv. Kingston. Differences in shoot to storage root ratiomay thus be associated with characteristics of the fibrous roots.Partition and specific activities are discussed in relationto the initiation and development of the storage organ. Key words: Daucus carota, carrot, assimilate, partition, ¹⁴C, storage root     

Effects of a Range of O2 Concentrations on Porosity of Barley Roots and on their Sugar and Protein Concentrations

Barley was grown at a range of oxygen concentrations (0.5–9mg l^–1), in nutrient solutions. Growth of both shootsand seminal roots was restricted by O₂ concentrations lowerthan 2–3 mg l^–1) but nodal root growth was not. Root porosities were increased even at those O₂ concentrationswhich did not restrict growth, and were inversely proportionalto the protein levels of the roots. Sugar concentrations increasedappreciably only at those O₂ concentrations which also restrictedgrowth. Hordeum vulgare L., barley, root porosity, sugar, protein, oxygen concentration     

Patterns of Assimilate Distribution and Source--sink Relationships in the Young Reproductive Tomato Plant (Lycopersicon esculentum Mill.)

Patterns of distribution of ¹⁴C were determined in 47-day-oldtomato plants (Lycopersicon esculentum Mill.) 24 h after theapplication of [¹⁴C]sucrose to individual source leaves fromleaves 1–10 (leaf 1 being the first leaf produced abovethe cotyledons). The first inflorescence of these plants wasbetween the ‘buds visible’ and the ‘firstanthesis’ stages of development. The predominant sink organs in these plants were the root system,the stem, the developing first inflorescence and the shoot ‘apex’(all tissues above node 10). The contribution made by individualsource leaves to the assimilate reaching these organs dependedupon the vertical position of the leaf on the main-stem axisand upon its position with respect to the phyllotactic arrangementof the leaves about this axis. The root system received assimilateprincipally from leaf 5 and higher leaves, and the stem apexfrom the four lowest leaves. The developing first inflorescencereceived assimilates mainly from leaves in the two orthostichiesadjacent to the radial position of the inflorescence on thevertical axis of the plant; these included leaves which weremajor contributors of ¹⁴C to the root system (leaves 6 and 8)and to the shoot apex (leaves 1 and 3). This pattern of distributionof assimilate may explain why root-restriction treatments andremoval of young leaves at the shoot apex can reduce the extentof flower bud abortion in the first inflorescence under conditionsof reduced photoassimilate availability. Lycopersicon esculentum Mill, tomato, assimilate distribution, source-sink relationships     

Effect of Rate of Photosynthesis on the Pattern of Assimilate Distribution in the Graminaceous Plant

Infrared gas analysis and a quantitative radiocarbon tracertechnique were used to measure photosynthesis and the distributionof ¹⁴C-labelled assimilate in Lolium temulentum and a uniculmbarley exposed continuously or intermittently to contrastinglight intensities. Plants grown for 10–20 d in low light(<50 W m^–2 of visible light) exported a greater proportionof their assimilate to growing leaves at the terminal meristemand a smaller proportion to their roots and tillers than equivalentplants in high light (150 W m^–2). A single day's exposure(8.4 h photoperiod) to a contrasting light regime elicited achange in the pattern of assimilate distribution in the samediurnal period, but 2–3 d exposure was required for asubstantial shift in the pattern of supply of assimilate tomeristems. The data indicated that in terms of assimilate distributioncomplete adaption to a new light regime is attained in about7 d.     

Root Morphology and Nitrogen Uptake of Maize Simultaneously Supplied with Ammonium and Nitrate in a Split-root System

We studied the response of maize (Zea mays L. cv. Anjou 256)to a simultaneous, but separated supply of ammonium and nitrate(localized supply, LS). A split-root system was used to supplyhalf of the roots with ammonium and the other half with nitrate.A homogeneously distributed supply of both nitrogen forms (HS)was the control treatment. Seedlings were grown for 12 d fromthe two-leaf to the three-leaf stage in hydroponics at threepH levels (4, 5·5 and 7). The total N concentration was3 mol m^-3. The split-root system was established by removingthe seminal root system and using only four nodal roots perplant. Total root length and root surface area were recordedautomatically with a modified Delta- T area meter. Other morphologicalroot traits (such as main axis length and diameter, number,density, and length of laterals) were recorded manually. Uptakeof ammonium and nitrate was measured by the depletion of thenutrient solution. As compared with LS, HS was superior in shootand root DM, total root length and root surface area, ammoniumand nitrate uptake and shoot nitrogen concentration, irrespectiveof pH level. This indicates that, also under field conditions,mixed ammonium and nitrate fertilization is only beneficialto plant growth if both N forms are evenly distributed in thesoil. At both HS and LS, ascending pH increased the ammonium:nitrateuptake ratio. At LS, declining pH induced a considerable shiftin the distribution of root DM, root length, and root surfacearea the nitrate-fed compartment.Copyright 1993, 1999 AcademicPress Maize, Zea may L., ammonium, nitrate, pH, root morphology, split-root     

Assimilation of 14CO2, Assimilate Translocation and Grain Yield in Three Primitive Nepalese Varieties and a European Cultivar (Senta) of Six-row Barley (Hordeum vulgare L.)

Three primitive varieties of six-row barley from Nepal withdifferent degrees of awn development were compared with a bredEuropean cultivar, Senta, under glasshouse conditions, fromanthesis onwards. Apart from the flag leaf the last three mainshoot leaves of the Nepalese varieties were smaller than thoseof Senta, and the lower leaves did not become senescent duringthe first 3 weeks of grain filling. In all varieties the penultimateleaf was the major assimilatory organ but in Senta the relativecontribution of the awns increased with time as the leaves senescedand fixed almost half of the total ¹⁴C assimilated by the shoot.Whereas the supply of assimilate to the ear increased with timein Senta, the reverse was found for the Nepalese varieties andoverall a smaller proportion of assimilate was supplied to theear in these varieties. The grain yield of the main shoot earof the two Nepalese varieties studied in detail was one-thirdof that of Senta; there were only approximately half the numberof grains in each median and lateral row of the ear and thegrains were smaller. There was no correlation between the degreeof awn development or the pattern of awn arrangement withinthe ear and the size of individual grains. The total biomassof the Nepalese varieties was much lower than that of Sentabut there were no differences in the harvest index or in tillerproduction. However, the Nepalese varieties, in contrast toSenta, had a mean grain yield per tiller comparable to thatof their main shoot and thus it appears that selection by plantbreeding may have increased the dominance of the main shoot. Hordeum vulgare L., barley, primitive barley varieties, carbon dioxide assimilation, assimilate translocation, grain filling     

The Influence of Plant Water Deficit on Distribution of 14C-labelled Assimilates in Cacao Seedlings

The relationship between plant water status and distributionof ¹⁴C-labelled assimilates in cacao (Theobroma cacao L.) wasevaluated after ¹⁴CO₂ pulse labelling leaves of seedlings subjectedto varying levels of water deficiency. The proportion of ¹⁴Cexported by source leaves was strongly affected by seedlingwater status. An increasing proportion of labelled assimilatesremained in source leaves at both 24-h and 72-h harvests aswater stress intensity increased. Water stress reduced the distributionof exported label to leaves and to the expanding flush in particularbut increased the proportion of label in stems and roots. Theresults suggest that current photoassimilates may be temporarilystored in source leaves and stems of cacao seedlings duringperiods of plant water deficit. The stress-induced changes inpartitioning of labelled carbon were in concordance with changesin shoot to root biomass ratios, which was likely due to greaterreduction in growth of above-ground organs to that of roots. Theobroma cacao L, assimilate partitioning, cacao, ¹⁴C-photoassimilate, water stress, water potential     

Effects of Plant Growth Regulators and Nutrient Supply on Tiller Bud Outgrowth in Barley (Hordeum distichum L.)

Seedlings of spring barley were raised in 100 and 20% nutrientsolution and treated with a foliar application of Terpal, Cerone,TIBA, GA₃ or BAP. The growth of individual tiller buds and tillers,the main shoot and the root system was recorded over the following15 d. Terpal and Cerone stimulated tiller bud elongation within5 d at both nutrient levels and after 15 d the number of emergedtillers was increased at the higher nutrient level. Terpal characteristicallypromoted the growth of secondary tiller buds whereas Ceronepromoted the emergence of the coleoptile tiller; both thesePGRs also retarded the development of the main shoot. TIBA increasedthe number of elongating tiller buds but this did not resultin greater tillering. GA₃ reduced the number of elongating tillerbuds and restricted their growth, especially in the high nutrientregime; this was accompanied by an increase in main shoot elongation.The growth and development of tiller buds was reduced by BAPand the number of emerged tillers was reduced at 15 d in bothnutrient levels; main shoot dry weight and root elongation werealso reduced. The results are considered in relation to theoverall influence of hormonal factors and mineral supply ontiller bud outgrowth. Hordeum distichum, spring barley, tiller bud outgrowth, plant growth regulators, Terpal, Cerone, GA₃, BAP, nutrient supply, apical dominance, TIBA     

Effect of Gibberellic Acid on the Distribution of Products of Photosynthesis in Sunflower

Single leaves on growing sunflower plants were allowed to assimilate¹⁴CO₂. Gibberellic acid was applied to the same leaf or to theterminal bud or the roots, and the distribution of assimilated¹⁴C was determined at intervals of 1–96 h. Gibberellicacid had no significant effect on initial distribution of ¹⁴Cduring the period of rapid export from the leaf, but enhancedre-export from the roots after translocation from the leaf hadvirtually ceased. Most of the ¹⁴C exported from the roots accumulatedin the shoot tip. The site of application of the hormone wasof relatively minor importance. Wherever it was applied themajor effect was enhancement of movement from the roots to theshoot tip. Application to the terminal bud was most effectivein this respect. There was no evidence that gibberellic aciddirectly affected the transport system, but the data supportthe hypothesis that it increases the strength of the sink inthe shoot tip. Helianthus annuus L., sunflower, assimilate transport, gibberellic acid, phloem transport     

Relationship Between the Development and Growth of Rye (Secale cereale L.) and the Potassium Concentration in Solution

The development and growth of rye (Secale cereale L. cv. Rheidol)was studied in seedlings grown hydroponically in complete nutrientsolutions containing between 10 and 600 µM K⁺. The phyllochron(defined as the interval between the appearance of successiveleaves) was used as a developmental timescale to compare plants.The pattern of both shoot and root development was strictlyordered on a phyllochron basis and was unaffected by solutionK⁺ concentration, with the exception that tillers in plantsgrown at the lowest K⁺ concentrations were occasionally eithernot initiated or aborted at an early stage of development. However,both the rate of leaf appearance on the main stem and successivetillers and the rate of tiller appearance were slower in plantsgrown at lower K⁺ concentrations. The rate of leaf appearanceon the main stem was reduced to below 90% of its maximal valueat solution concentrations below about 50 µM K⁺. Plantrelative growth rate (RGR) was also reduced by lowering theK⁺ concentration of the nutrient solution and fell to below90% of its maximal value at solution concentrations below about200 µM K⁺. There was a complex relationship between tissueK⁺ concentration and the K⁺ concentration of the nutrient solution,which differed between leaves and root. Leaf K⁺ concentrationincreased steadily from about 50 µmol g^-1 f. wt to about200 µmol g^-1 f. wt as solution K⁺ concentration was increasedfrom 10 to 400 µM. In contrast, root K⁺ concentrationexhibited a sigmoidal dependence on solution K⁺ concentration,maintaining a minimal value of approximately 20 µmol g^-1f. wt at concentration below 100 µM K⁺, then increasingprogressively to about 120 µmol g^-1 f. wt at a solutionconcentration of 600 µM K⁺. The 'critical' leaf K⁺ concentration,i.e. the concentration at which either plant RGR or plant developmentwas reduced 90% of its maximal value, was 86 µmol g^-1f. wt for plant RGR and 150 µmol g^-1 f. wt for plant development.The 'critical' root K⁺ concentration was 24 µmol g^-1 f.wt K⁺ for both RGR and development. A decline in tissue K⁺ concentrationbelow these thresholds reduced plant growth considerably. RootK⁺ concentration was a sensitive indicator of the K⁺ statusof the plant with respect to potential growth since plant growthdeclined abruptly as root K⁺ concentration approached its 'critical'value, whereas plant growth showed a less defined relationshipwith shoot K⁺ concentration.Copyright 1993, 1999 Academic Press Critical K⁺ concentration, development, potassium, relative growth rate (RGR), rye, Secale cereale L. cv. Rheidol     

Vegetable Plant Part Relationships. III. Modification of Carrot (Daucus carota L.) Root and Shoot Weights by Gibberellic Acid and Daminozide

Aqueous foliar sprays of N-dimethylaminosuccinamic acid (daminozide)at 2000 p.p.m. and gibberellic acid (GA) at 100 p.p.m. wereapplied 45, 59, 82 and 100 days after sowing to Chantenay carrotswith population densities of 244, 495 and 883 plants m^–2.The plants were harvested on ten approximately weekly occasions;fresh weights were determined and d. wt estimates were obtainedfor the separated shoots (s) and roots (r). Allometric linearregressions of the logarithm of s on that of r at each harvestseparately, clearly showed that GA always increased shoot: rootratio and reduced root yield (by approximately 35 per cent)but could sometimes also increase whole-plant weight. Daminozideincreased root yield (by approximately 7 per cent from 80 tonnesha^–1) and tended to have effects opposite to those ofGA. Daucus carota L., carrot, root weight, shoot weight, N-dimethylaminosuccinamic acid (daminozide), gibberellic acid     

Growth of Zea mays L. plants with their seminal roots only. Effects on plant development, xylem transport, mineral nutrition and the flow and distribution of abscisic acid (ABA) as a possible shoot to root signal

Maize (Zea mays L.) was grown in quartz sand culture eitherwith a normal root system (controls) or with seminal roots only(‘single-rooted’). Development of adventitious rootswas prevented by using plants with an etiolated mesocotyl andthe stem base was positioned 5–8 cm above the sand. Eventhough the roots of the single-rooted plants were sufficientlysupplied with water and nutrients, the leaves experienced waterdeficits and showed decreased transpiration as trans plrationalwater flow was restricted by the constant number of xylem vesselspresent in the mesocotyl. As a consequence of this restriction,transpirational water flow velocities in the metaxylem vesselsreached mean values of 270 m h^–1 and phloem transportvelocities of 5.2 m h^–1. Despite limited xylem transportmineral nutrient concentrations in leaf tissues were not decreasedin single-rooted plants, but shoot and particularly stem developmentwas somewhat inhibited. Due to the lack of adventitious rootsthe shoot:root ratio was strongly increased in the single-rootedplants, but the seminal roots showed compensatory growth comparedto those in control plants. Consistent with decreased leaf conductance,ABA concentrations in leaves of single-rooted plants were elevatedup to 10-fold, but xylem sap ABA concentrations in these plantswere lower than in controls, in good agreement with the well-wateredconditions experienced by the seminal roots. Surprisingly, however,ABA concentrations in tissues of the seminal roots of the single-rooted plants were clearly increased compared to the controls,presumably due to increased ABA import via phloem from the water-stressedleaves. The results are discussed in relation to the role ofABA as a shoot to root signal. Key words: Zea mays, seminal roots, plant development, xylem transport, mineral nutrition, ABA, shoot-to-root signal     

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.