The Influence of Leaf Position and Defoliation on the Assimilation and Translocation of Carbon in White Clover (Trifolium repens L.) 1. Carbon Distribution Patterns

The assimilation of carbon (C) by, and distribution of ¹⁴C from,leaves at each end of an unbroken sequence of ten mature leaveson the main stolon of clonal plants of white clover (Trifoliumrepens L.) were measured to identify intra-plant factors determiningthe direction of C movement from leaves. Leaves at two intermediatepositions were also measured. Localized movement of ¹⁴C to sinks at the same node as, or atthe one to two nodes immediately behind, the fed leaf accountedfor 40–50% of the total ¹⁴C exported by all measured leaves.A further 50–60% of exported ¹⁴C was therefore availablefor more-distant sinks, and the direction of translocation ofthis C was determined by the relative total strength or demand(number x size x rate of activity or growth) of sinks forwardof, or behind, the leaf in question. Thus 85% of the ¹⁴C exportedfrom the youngest measured leaf moved toward the base of thestolon, while about 60% of the ¹⁴C exported from the oldestleaf moved acropetally. Defoliating plants to leave just one mature leaf on the mainstolon (at any one of the same four positions studied in undefoliatedplants), and no leaves on branches, resulted in: (1) increasednet photosynthetic rate in all residual leaves: (2) increased%export of fixed C from one of the four leaves; (3) increasedexport to the main stolon apex from all except the eldest leaf;(4) increased export to branches from three of the four leaves;and (5) decreased export to stolon tissue and roots from allleaves, within 3 d of defoliation. These responses would seemto ensure the fastest possible replacement of lost leaf areaand, thus, restoration of homeostatic growth. The observed patternsof C assimilation and distribution in both undefoliated anddefoliated white clover plants are consistent with the generalrules of source-sink theory; the distance between sources andcompeting sinks, and relative sink strength, emerge as the mostimportant intra-plant factors governing C movement. These resultsemphasize the need to consider plant morphology, and the modularnature of plant growth, when interpreting patterns of resourceallocation in clonal plants, or plant responses to stressessuch as partial defoliation. Trifolium repens L, white clover, photosynthesis, assimilate translocation, defoliation     

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.     

Pattern of defoliation and its effect on photosynthesis and growth of Goldenrod

1. Leaf area was removed from Solidago altissima in either a dispersed pattern (half of every leaf removed) or a concentrated pattern (every other leaf removed) and effects on leaf gas exchange, vegetative growth and flowering were examined relative to undefoliated controls. Gas exchange was measured for leaves remaining after defoliation and for regrowth leaves that developed post-damage (at 7, 16 and 26 days post-defoliation).
2. Area-based photosynthetic rates of leaves remaining after defoliation were not affected by either dispersed or concentrated damage, but damage of both types enhanced area-based photosynthesis of regrowth leaves at 16 days post-defoliation and to a lesser extent at 26 days post-defoliation.
3. Dispersed damage, but not concentrated damage, stimulated mass-based photosynthesis of undamaged leaves remaining after defoliation. Undamaged leaves remaining after defoliation and regrowth leaves on damaged plants had higher specific leaf area (leaf area/leaf mass) than comparable leaves on control plants. Mass-based photosynthesis was more strongly elevated by defoliation than area-based photosynthesis because of this increase in specific leaf area.
4. Plants with dispersed damage recovered more quickly from defoliation; they had higher relative growth rates in the first week post-defoliation than plants with concentrated damage. Both types of defoliation caused similar reductions in flower production.
5. These results add to accumulating evidence that dispersed damage is generally less detrimental to plants than concentrated damage and suggest that physiological changes in leaves may be part of the reason.     

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.     

Defoliation in White Clover: Regrowth, Photosynthesis and N2 Fixation

Single plants of white clover, grown in a controlled environmentand dependent for nitrogen on fixation in their root nodules,were defoliated once by removing approximately half their shoottissue. Their regrowth was compared with the growth of comparableundefoliated plants. Two similar experiments were carried out:in the first, plants were defoliated at 2.5 g, and in the secondat 1.2 g total plant d. wt. Defoliation reduced rate of N₂ fixation by > 70 per cent,rate of photosynthesis by 83–96 per cent, and rate ofplant respiration by 30–40 per cent. Nodule weights initiallydeclined following defoliation as a result of loss of carbohydratesand other unidentified components. No immediate shedding ofnodules was observed but nodules on the most severely defoliatedplants exhibited accelerated senescence. The original rates of N₂ fixation were re-attained after 5–6or 9 d regrowth, with increase in plant size at defoliation.In general, the rate of recovery of N₂ fixation was relatedto the re-establishment and increase of the plant's photosyntheticcapacity. Throughout the growth of both defoliated and undefoliatedplants nodule respiration (metabolism) accounted for at least23 ± 2 per cent of gross photosynthesis. The unit ‘cost’of fixing N₂ in root nodules, in terms of photosynthate, appearedto be unaffected by defoliation, except perhaps for plants veryrecently defoliated. Similarly, the percentage nitrogen contentsof shoot, root and nodules of defoliated plants became adaptedwithin a few days to those characteristic of undefoliated plants. Trifolium repens, white clover, N₂ fixation, defoliation, photosynthesis, respiration     

The utilization of recently assimilated carbon in graminaceous plants

Isotopic carbon and infra-red gas analysis techniques were used to measure the following growth attributes in maize, sorghum, winter wheat and perennial ryegrass: the rate of entry of carbon into each main shoot leaf; the rate of translocation of leaf assimilate to meristems; the fraction of leaf and total shoot assimilate respired in one diurnal period; and the distribution of residual assimilate to new leaf, stem, axillary shoots and root. The two tropical plants possessed higher leaf assimilation rates and larger leaves than the temperate species, but their efficiency of translocation was only marginally superior in the experimental conditions. In all species, c. 25% of the assimilate generated in the 8·4h photoperiod was respired in in the same diurnal period. Maize and sorghum partitioned a greater proportion of their total shoot assimilate to new leaf tissue at the main shoot apex and to root than wheat and ryegrass. On the other hand, wheat and ryegrass exported up to 30% of their assimilate to axillary shoots; in sorghum, little assimilate was translocated to axillary shoots, while in maize this activity was completely absent. Plant habit, as exemplified by the contrast between the annual, single-axis maize plant and the perennial, multi-tillering ryegrass plant, appears to be a reflexion of the pattern of assimilate distribution to areas of potential growth. With the exception of superior leaf assimilation rates in maize and sorghum, the four species showed no marked differences in respect of the production, transport and respiratory utilization of assimilates.     

Combined Effects of Partial Defoliation and Nutrient Availability on Cloned Betula pendula Saplings: II. CHANGES IN NET PHOTOSYNTHESIS AND RELATED BIOCHEMICAL PROPERTIES

The combined effects of partial defoliation and nutrient availabilityon net photosynthesis and related biochemical variables werestudied in cloned Betula pendula Roth saplings. The saplingswere randomly assigned to different nutrient levels (5, 1·5and 0·5 mol N m^–3) in aerated nutrient cultureand to the following defoliation treatments: (1) control (nodamage), (2) damage of the developing main stem leaves (halfof the leaf lamina removed), and (3)removal of the developingmain stem leaves (entire leaf lamina removed). The leaf immediatelybelow the damaged area in the treated plants, and the correspondingleaf in the control plants, were selected for study. Net photosynthesismeasurements and biochemical determinations were made 2, 8 and14 d after assigning the treatments. At intermediate and lownutrient levels the final net photosynthetic capacity was significantlyhigher in the saplings with the topmost leaves removed thanin the undamaged control saplings, indicating that the expressionof compensatory photosynthesis after partial defoliation isnot inhibited by nutrient deficiency. The photosynthetic enhancementwas closely associated with the increased initial activity ofribulose 1,5-bisphosphate carboxylase/oxygenase (Rubisco). However,the increased activity of Rubisco was not exclusively the resultof a higher amount of Rubisco. The expression of compensatoryphotosynthesis after partial defoliation in our study cannotunequivocally be attributed to an increased flow of nitrogento the remaining leaves. Key words: Partial defoliation, nutrient availability, net photosynthesis, nitrogen, Rubisco     

The Export and Distribution of 14C-labelled Assimilates from each Leaf on the Shoot of Lolium temulentum during Reproductive and Vegetative Growth

In both reproductive and vegetative plants of Lolium temulentumL., the export of ¹⁴C-labelled assimilates from each healthyleaf on the main shoot to terminal meristem, stem, tillers,and roots was measured each time a new leaf was expanded, fora period of 5 to 6 weeks. Some labelled assimilates moved fromeach leaf on the main shoot to every meristem in the same shoot,as well as to the tops and roots of adjacent organically attachedtillers. The terminal meristem of the reproductive shoot, which includedthe developing inflorescence, received 70–80 per centof the carbon assimilated by the emerged portion of the growingleaf, 15–25 per cent of the carbon assimilated by thetwo youngest expanded leaves, and 5–10 per cent of thatfrom each of the older leaves. A similar pattern of carbon supplyto the terminal meristem was found in vegetative shoots, exceptthat older leaves on young vegetative shoots supplied even lessof their carbon to the terminal meristem. The general conclusionis that developing leaves at the tip of the shoot receive aboutthe same proportion of carbon from each leaf as does a developinginflorescence. Young expanded leaves provided most labelled assimilates forstem growth; during both reproductive and vegetative growth,expanded leaves increased their export of labelled carbon tostem, and exported less of their ¹⁴C to roots and sometimesto tillers. In these reproductive and vegetative shoots, grown in a constantexternal environment, the major changes in the pattern of distributionof labelled assimilates appeared to be the result of increasedmeristematic activity in stem internodes; the development ofan inflorescence had no obvious direct effect on the carboneconomy of shoots.     

Long-term Partitioning, Storage and Remobilization of 14C Assimilated by Trifolium repens (cv. Blanca)

The fourth fully expanded leaf on the main stolon of white cloverplants was exposed to ¹⁴CO₂. Thereafter, quantitative and fractionalanalysis of the partitioning, storage and remobilization afterdefoliation of the ¹⁴C labelled assimilate was sequentiallyconducted over a 2- to 3-week period. In undefoliated plants, most ¹⁴C reached its final destinationwithin 24 h of feeding. Forty percent of assimilated ¹⁴C waslost through respiration, while the rest was exported, predominantlyto meristems, but also to roots, stolons and leaves. The ¹⁴Cinitially translocated to meristems was subsequently recoveredin stolon and leaf tissue as the plants matured. Approximately 10% of assimilated ¹⁴C was invested into long-termstorage in roots and stolons. These reserves were remobilizedafter both partial and total defoliation, and a portion of theremobilized ¹⁴C was incorporated into new growth, Partly defoliatedplants regrew more rapidly than totally defoliated plants, butmore ¹⁴C reserve depletion took place in the totally defoliatedtreatment. Reserve depletion took place from both stolons androots, but stolon reserves were preferentially utilized. Bothhigh and low molecular weight storage compounds were involved. Trifolium repens, white clover, assimilate partitioning, storage, remobilization, defoliation     

Sectorial root growth in cuttings of Coleus rehneltianus in response to localized aerial defoliation

Asymmetries in root growth in response to localized aerial defoliation were examined in Coleus rehneltianus (Lamiaceae). We confirmed that assimilate transport was sectorial by examining the distribution of ¹⁴C-labeled carbohydrates following a 24-h chase period. Integrated physiological units (IPUs), or sectors, extended from the leaves into the roots, and this was reflected in the differential growth of roots following artificial defoliation of part of the leaf canopy. When defoliation was localized within leaves or leaf halves within sectors, roots grew asymmetrically, with decreased root growth in defoliated sectors. Three root populations were identified by their location and growth responses: stem side, stem corner, and bottom side roots, and asymmetric growth was observed in all three populations. Only the growth of stem corner roots, which made up 35–90% of dry mass of the total root population, was influenced by the pattern of aerial defoliation. In contrast, asymmetries in the growth of the other two root populations appeared to reflect the distribution of leaf biomass prior to defoliation.     

Regrowth of spring canola (Brassica napus) after defoliation

Aims

Regrowth of dual-purpose canola after grazing is important for commercial success and the aim of this research was to investigate the effects of defoliation on the development, growth, photosynthesis and allocation of carbohydrates.

Methods

We conducted two pot experiments in which defoliation was conducted at multiple intensities with scissors. Experiment 1 determined changes in flowering date due to defoliation while Experiment 2 investigated the effects of defoliation on growth, photosynthesis and allocation of carbohydrates in canola.

Results

Time to the appearance of the first flower was delayed by up to 9 days after the removal of all leaves at the start of stem elongation (GS30), and up to 19 days if the elongating bud was also removed. Stem growth rate decreased by 56–86 % due to defoliation and tap roots did not increase in mass when plants were completely defoliated. Leaf area continued to expand at the same rate as in un-defoliated plants. The new leaf area established per gram of regrowth biomass over 20 days was 158 cm².g^-1 for the complete defoliation treatments compared with 27 cm².g^?1 for the half-defoliated treatment and 13 cm².g^?1 for the un-defoliated treatment. Despite a reduction in total biomass of up to 60 %, the proportion of dry matter partitioned to the leaves was 18 % for all treatments within 20 days after defoliation. Total non-structural carbohydrate levels were reduced rapidly in the stem by day two (predominately sucrose) and the tap root by day four (predominately starch) after defoliation and did not recover to match un-defoliated plant levels within 20 days. Residual leaves on defoliated plants maintained photosynthetic rate compared with the same leaf cohorts on un-defoliated plants in which photosynthetic rate decreased to 39 % by day 12.

Conclusions

The rapid recovery of leaf area in defoliated canola was facilitated by the sustained high photosynthetic rate in remaining leaves, rapid mobilisation of stored sugars (stem) and starch (root), and a cessation of root and stem growth.     

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     

Partitioning of reserve and newly assimilated carbon in roots and leaf tissues of Lolium perenne during regrowth after defoliation: assessment by 13C steady-state labelling and carbohydrate analysis

The relative significance of the use of stored or currently absorbed C for the growth of leaves or roots of Lolium perenne L. after defoliation was assessed by steady-state labelling of atmospheric CO₂. Leaf growth for the first two days after defoliation was to a large extent dependent on the use of C reserves. The basal part of the elongating leaves was mainly new tissue and 91% of the C in this part of the leaf was derived from reserves assimilated prior to defoliation. However, half of the sucrose in the growth zone was produced from photosynthesis by the emerged leaves. Fructans that were initially present in elongating leaf bases were hydrolysed (loss of 93 to 100%) and the resulting fructose was found in the new leaf bases, suggesting that this pool may be used to support cell division and elongation. Despite a negative C balance at the whole-plant level, fructans were synthesized from sucrose that was translocated to the new leaf bases. After a regrowth period of 28 d, 45% of the C fixed before defoliation was still present in the root and leaf tissue and only 1% was incorporated in entirely new tissue.     

Plant-herbivore interactions: Examination of potential effects of bison saliva on regrowth of Bouteloua gracilis (H.B.K.) lag

Summary Laboratory experiments were performed to determine whether regrowth of blue grama was affected by potential growth-promoting substances in saliva of North American bison. We observed no statistically significant effects of foliar application of whole bison saliva on net photosynthesis (P_N), root respiration (R_R), allocation patterns of photosynthetically fixed ¹⁴C, or regrowth rates over a 10-day period following clipping to various heights. In a 10-week experiment, there were no significant effects of saliva on leaf, crown or root growth or tiller production in plants clipped to heights of 6, 4 or 2 cm above crowns. Similarly, nitrogen-stressed plants failed to show significant changes in growth rates or tillering in response to saliva over a 3-week period. Clipped blue grama plants did exhibit significant compensatory growth responses, including higher P_N rates from 3–10 days following clipping and allocation of a higher proportion of current photosynthate to synthesis of new leaf tissue with increasing severity of defoliation. Nevertheless, unclipped plants invariably outproduced clipped plants following defoliation.     

Root Development and Source-Sink Relations in Carrot, Daucus carota L: II. EFFECTS OF ROOT PRUNING ON CARBON ASSIMILATION AND THE PARTITIONING OF ASSIMILATES

Physiological responses to root pruning were investigated bycomparing ¹⁴CO₂ fixation rates, the partitioning of ¹⁴C-labelledassimilate, and soluble and insoluble carbohydrate levels inthe leaves of carrot plants following the removal of some ofthe fibrous roots, or fibrous roots and part of the tap root.Root pruning reduced ¹⁴CO₂ fixation by 28–45% but leafspecific activity (¹⁴C assimilation g-¹ leaf fresh weight) wasunchanged. The proportion of total assimilate exported to theroot system increased following root pruning and this was atthe expense of the developing leaves. In younger plants (wherethe tap root received 10% of the assimilate) the supply of ¹⁴Cto the tap root was maintained in spite of root pruning. However,shortening the tap root to 3 cm in older plants (in which 30%of the fixed 14C was normally exported to the developing storageorgan), reduced its sink capacity and resulted in slightly greaterretention of ¹⁴C in the mature leaves. Greater concentrationsof insoluble carbohydrate were found in the mature leaves followingroot pruning but soluble sugar content was unaffected. Onlysmall differences were observed in the distribution of ¹⁴C betweensoluble and insoluble carbohydrate fractions when plants werefed ¹⁴CO₂ several days after the root pruning operations. Thesephysiological responses were mainly associated with the removalof fibrous roots and support the view that the fibrous rootsystem is more important than the developing storage organ inregulating growth in young carrot plants.     

Carbon Partitioning in Mature Leaves of Pepper: Effects of Daylength

Grange, R. 1. 1985. Carbon partitioning in mature leaves ofpepper: effects of daylength.—J. exp. Bot. 36: 1749–1759. The partitioning of recently fixed carbon has been examinedin mature pepper leaves grown in 6, 10 or 14 h photoperiodsat different irradiances chosen to give similar radiation integralsand in a 6 h photoperiod at the lowest of these irradiances.The partitioning of carbon into export, starch, sugars and respirationwas followed over the photopenod and the subsequent night ina mature leaf. The maximum export rate during the day (approximately 18 µgC cm^–2 leaf h^–1) was not significantly differentamong the treatments. Net photosynthesis rate was directly relatedto irradiance; the proportion of net photosynthesis exportedduring the day was 33% in 6-h days and 57% in 14-h days. Leafstarch accumulation (as a proportion of net photosynthesis rate)increased slightly when plants were grown in 6-h days. The remobilization of starch and sugars at night allowed exportrates to remain similar over 24 h when plants were grown in10-h or 14-h photoperiods. Leaves grown in 6-h days showed nosignificant changes in export rate during the first few hoursof night but exhausted their starch reserves during the nightand export rates declined. Sucrose and hexose levels decreased at the onset of darkness,but did not fall below 40 µg cm^–2 in plants grownin 10-h or 14-h photoperiods; when this level was reached after3–4 h of darkness, starch breakdown began. In leaves grownin both 6-h treatments, sucrose levels fell below 40 µgcm^–2 when starch reserves were depleted during the nightand the export rate decreased concurrently. The results are discussed in relation to the control of exportand starch metabolism in the leaf. Key words: Pepper, partitioning, daylength     

Defoliation,Photosynthetic Rates,and Assimilate Transport in Grapevine Plants

The source-sink relations in grapevine (Vitis vinifera L., var. Rkatsiteli) plants were disturbed by defoliation at different stages of vegetative growth in order to investigate changes in photosynthetic activity and assimilate partitioning. Defoliation was shown to stimulate photosynthesis in the remaining source leaves, enhance the assimilate export, and diminish the midday suppression of photosynthesis. Defoliation created a powerful sink for assimilates, and stimulated their delivery to the affected zone. It is hypothesized that defoliation-induced stress is accompanied by a substantial enhancement of photosynthetic activity and by redistribution of assimilate flows, which enables a sustained supply of assimilates to the sink organs of grapevine plants.__________Translated from Fiziologiya Rastenii, Vol. 52, No. 4, 2005, pp. 507–512.Original Russian Text Copyright © 2005 by Chanishvili, Badridze, Barblishvili, Dolidze.     

Roles of the fructans from leaf sheaths and from the elongating leaf bases in the regrowth following defoliation of Lolium perenne L

The study of carbohydrate metabolism in perennial ryegrass (Lolium perenne L. cv. Bravo) during the first 48 h of regrowth showed that fructans from elongating leaf bases were hydrolysed first whereas fructans in mature leaf sheaths were degraded only after a lag of 1.5 h. In elongating leaf bases, the decline in fructan content occurred not only in the differentiation zone (30–60 mm from the leaf base), but also in the growth zone. Unlike other soluble carbohydrates, the net deposition rate of fructose remained positive and even rose during the first day following defoliation. The activity of fructan exohydrolase (FEH; EC 3.2.1.80) was maximal in the differentiation zone before defoliation and increased in all segments, but peaked in the growth zone after defoliation. These data strongly indicate that fructans stored in the leaf growth zone were hydrolysed and recycled in that zone to sustain the refoliation immediately after defoliation. Despite the depletion of carbohydrates, leaves of defoliated plants elongated at a significantly higher rate than those of undefoliated plants, during the first 10 h of regrowth. This can be partly attributed to the transient increase in water and nitrate deposition rate. The results are discussed in relation to defoliation tolerance. Received: 16 June 2000 / Accepted: 17 October 2000     

Regulation of Assimilate Translocation Between Leaves and Fruits in the Tomato

Simultaneous measurement of export from leaves and import tofruits were made on tomato plants reduced to one fully expandedleaf and one fruit. Experimental leaves were exposed to sixlight flux densities (0.5–100 W m^–2) for 24 h whilerapidly growing fruits were kept in the dark at 22 °C. The rates of export of assimilate from these leaves varied from70 to 120 mg C leaf^–1 day^–1 corresponding with ratesof carbon fixation from 3 to 290 mg C leaf^–1 day^–1.Export from leaves with the lowest carbon fixation rates weremaintained by a loss of up to one-sixth of their initial carbon.In contrast, leaves with the highest carbon fixation rates exportedonly half the newly fixed carbon. The rates of import of assimilate to similar-sized fruits (c.16 cm³) were between 80 and 110 mg C fr^–1 day^–1but differed from the export rates of the source leaves. Thespecific growth rates and the specific respiration rates ofthe fruits were related to their initial carbon content at thebeginning of the experiment. Thus, over 24 h, the rate of importwas predetermined by the developmental stage of the fruit unalteredby the rate of current carbon fixation in the source leaf. Translocationof assimilate was regulated by sink demand under both source-and sink-limiting conditions in this short-term situation. The dynamic relationship between assimilate production in leavesand its utilization in fruits is discussed together with therole of sucrose concentration in these organs in regulatingtransport. Lycopersicon esculentumL, tomato assimilate translocation, source-sink relationships     

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.