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     

Dry Matter Distribution between Shoot and Storage Root of Carrot, Parsnip, Radish and Red Beet

The relationship between shoot and storage root weights of carrot,parsnip, radish and red beet was examined using plants of differentsize and age. For both carrot and parsnip plants of the sameage there was a linear relationship between the logarithms ofshoot and storage root d. wts which was unaffected by plantdensity. For parsnip the parameters of the regression were unchangedby harvest time, while for carrot the slope of the regressionremained constant and the intercept progressively declined withharvest time. These observations are consistent with an hypothesisof assimilate partition based on competition between sinks.For both radish and red beet at very high densities the logarithmicrelationship between shoot and storage root weight was curvilinearwith significant effects of both harvest time and density. Itis suggested that interplant competition may affect the initiationof storage tissue development in both these species. It is concludedthat for radish and red beet, even when considering only plantsproducing normal swollen storage roots, partition of assimilateis affected by factors not incorporated into the above hypothesis. Root crop, carrot, parsnip, radish, red beet, partition of assimilated carbon, model     

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     

Vegetable Plant Part Relationships. IV. An Interpretation of Growth Regulator Experiments with Root Crops

A quantitative scheme to describe the growth of plant partsduring the vegetative phase of development is extended to includethe concept of hormone directed transport. The expected effecton shoot/storage root d. wt relationships of a single àxternalapplication of a growth regulating chemical is investigatedusing this approach. The theory predicts that the resultingchange in the pattern of assimilate partitioning will modifythe underlying relationship between shoot and storage root weightsin a simple measurable fashion. The theory is inevitably oversimplifiedand speculative in parts but when a derived equation was fittedto data from an experiment in which gibberellic acid and daminozidewere applied to carrots, close agreement was found between thepredictions and experimental data. Daucus carota, carrot, shoot weight, root weight, gibberellic acid, N-dimethylamino-succinamic acid (daminozide), growth analysis     

Vegetable Plant Part Relationships. II. A Quantitative Hypothesis for Shoot/Storage Root Development

A simple quantitative formulation of the concept of the controlof partitioning of assimilated carbon by the behaviour of plantcomponents as competing sinks is developed. An equation, In s = + In r—t, relating shoot (s) and storage root (r) d. wts, and the lengthof growth period (t), is constructed by considering possiblefates of imported assimilates into different plant parts. Thevalues of the equations' parameters depend on the relative sinkactivities of the plant parts, tissue respiration rates andinitial weights of plant components. The equation closely fitteddata collected from a number of carrot and beet experimentsin which planting density had been varied. Estimates of shootand storage root maintenance respiration rates, derived fromthe parameter , were of the correct order of magnitude. Othersets of experimental data are also discussed in the light ofpredictions of the theory and possible uses and extensions ofthis approach to assimilate partitioning are briefly discussed. Daucus carota L., Beta vulgaris, carrot, red beet, partition of assimilated carbon, maintenance respiration, storage root     

Assimilate Partitioning in Red and White Clover Either Dependent on N2 Fixation in Root Nodules or Utilizing Nitrate Nitrogen

During vegetative growth in controlled environments, the patternof distribution of ¹⁴C-labelled assimilates to shoot and root,and to the meristems of the shoot, was measured in red and whiteclover plants either wholly dependent on N₂ fixation in rootnodules or receiving abundant nitrate nitrogen but lacking nodules. In experiments where single leaves on the primary shoot wereexposed to ¹⁴CO₂, nodulated plants of both clovers generallyexported more of their labelled assimilates to root (+nodules),than equivalent plants utilizing nitrate nitrogen, and thiswas offset by reduced export to branches (red clover) or stolons(white clover). The intensity of these effects varied with experiment.The export of labelled assimilate to growing leaves at the terminalmeristem of the donor shoot was not influenced by source ofnitrogen. Internode elongation in the donor shoot utilized nolabelled assimilate. Whole plants of white clover exposed to ¹⁴CO₂ on seven occasionsover 32 days exhibited the same effect on export to root (+nodules),which increased slightly in intensity with increasing plantage. Nodulated plants had larger root: shoot ratios than theirequivalents utilizing nitrate nitrogen. Trifolium repens, Trifolium pratense, red clover, white clover, nitrogen fixation, nitrate utilization, assimilate partitioning     

Comparative Analysis of Tuber Development in Six Sweet Potato (Ipomoea batatas (L.) Lam) Cultivars: 1. Tuber initiation, tuber growth and partition of assimilate

Differences in tuber initiation, tuber growth and partitionof assimilate during development were investigated in six localsweet potato cultivars. Study of the quantitative morphogenesisof root types in the sweet potato root system indicated thattuber initiation was completed by 8 weeks after planting inmost cultivars and frustrated thereafter. Subsequent differencesin patterns of tuber development were not obviously relatedto final tuber yield. Thus, high yield resulted either froma short period of rapid tuber growth or a longer period of slowertuber growth. Maximum shoot growth was achieved by week 12 inall except one cultivar, and the cessation of shoot growth resultedin either increases or decreases in tuber growth rate or hadno affect on the rate of tuber growth (cv. A28/7). The partitionof assimilate to tubers at final harvest was the parameter mostclosely related to the yield of the six cultivars studied, butthere was some evidence that such partition was related to totaldry weight. It is suggested that limiting factors in sweet potatotuber yield, e.g. assimilate production and transport or capacityfor tuber growth varied with the cultivar studied.     

Role of Foliage Habit in the Competition Between Differently Sized Plants in Carrot Crops

The length of the tallest leaf on a plant varied considerablywithin each of three carrot crops, and in each crop there wasa high positive correlation between shoot height and root weight.Lowering the petiole angle of carrot leaves greatly reducedthe ability of plants to compete against untreated neighbours,but did not affect growth when all plants were treated similarly.Cutting treatments which produced a more uniform shoot heightfor all plants, allowed small plants temporarily to overcomethe dominance of their larger neighbours. Similar cutting treatmentsapplied in a field experiment reduced crop growth and variationin root weight. These findings support the view that shoot heightdifference is an important component of inter-plant competitionin carrot crops and hence is a source of root-weight variation. Daucus carota L., carrot, foliage habit, competition     

Dry Matter Distribution Between the Shoot and Storage Root of Carrot (Daucus carota L.): II. Effect of Foliar Application of Gibberellic Acid

The effects of foliar sprays of gibberellic acid (GA) on drymatter distribution in carrots were investigated in three fieldexperiments between 1977 and 1979. The results were interpretedby using a simple quantitative model of assimilate partitioningbetween storage root and shoot. Although the response to GAvaried depending on the year in which the experiment was done,the model successfully described the effects of spraying time,GA concentration, planting density, plant age and cultivar.The maximum effects on repartitioning were achieved with concentrationsof 100–500 mg 1_–1 GA and, except for very earlyor late sprays, the effect was constant with time over a roughly4-week period. The multiplicative nature of the model was usefulin that despite the occurrence of various interactions, a similarassimilate partitioning response to GA in all four cultivarswas demonstrated. The model did not suggest an explanation forthe seasonal effects. Daucus carota L., carrot, assimilate distribution, dry matter distribution, storage root, gibberellic acid, partitioning model     

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     

Seasonal Changes in the Physiology of S24 Perennial Ryegrass (Lolium perenne L.): 3. Partition of Assimilates between Root and Shoot during the Transition from Vegetative to Reproductive Growth

The partition of ¹⁴C labelled current assimilates to root insimulated swards of Lolium perenne cv. S24 was measured duringthe transition from vegetative growth in autumn to reproductivegrowth in spring under close to natural conditions of lightand temperature. Assimilate partitioning was also measured in‘established’ swards cut three times during thegrowing season and in vegetative ‘seedling’ swardsgrowing in autumn and in spring. All measurements were madewhen the swards had achieved more than 90 per cent light interception,and all swards were abundantly supplied with water and mineralnutrients. During autumn there was a gradual decrease in the proportionof assimilates partitioned to the roots in both the ‘established’and the ‘seedling’ swards. In the established swards,partition to roots was low over winter, increased during earlyspring, but decreased dramatically, later in the spring, whenstem elongation began. In contrast, in the unvernalized vegatativeseedling swards in spring, partition to roots remained high. The seasonal pattern of assimilate partitioning is consideredin relation to changes in the natural environment and the rateat which the crop fixed carbon in photosynthesis. A decreasein the proportion of assimilates partitioned to roots duringlate spring was significant in increasing the production ofshoot at that time but seasonal differences in partition contributedvery little to the marked differences in shoot growth betweenthe spring and autumn crop. Lolium perenne L., perennial ryegrass, partition of assimilates, flowering     

Effect of Assimilate Supply on Development and Growth Potential of Axillary Buds in Roses

The effect of assimilate supply on axillary bud developmentand subsequent shoot growth was investigated in roses. Differencesin assimilate supply were imposed by differential defoliation.Fresh and dry mass of axillary buds increased with increasedassimilate supply. The growth potential of buds was studiedeither by pruning the parent shoot above the bud, by graftingthe bud or by culturing the bud in vitro. Time until bud breakwas not clearly affected by assimilate supply during bud development,Increase in assimilate supply slightly increased the numberof leaves and leaf primordia in the bud; the number of leavespreceding the flower on the shoot grown from the axillary budsubstantially increased. No difference was found in the numberof leaves preceding the flower on shoots grown from buds attachedto the parent shoot and those from buds grafted on a cutting,indicating that at the moment of release from inhibition thebud meristem became determined to produce a specific numberof leaves and to develop into a flower. Assimilate supply duringaxillary bud development increased the number of pith cells,but the final size of the pith in the subsequent shoot was largelydetermined by cell enlargement, which was dependent on assimilatesupply during shoot growth. Shoot growth after release frominhibition was affected by assimilate supply during axillarybud development only when buds sprouted attached to the parentshoot, indicating that shoot growth is, to a major extent, dependenton the assimilate supply available while growth is taking place.Copyright1994, 1999 Academic Press Assimilate supply, axillary bud, cell number, cell size, defoliation, development, growth potential, meristem programming, pith, Rosa hybrida, rose, shoot growth     

Effects of gibberellic acid and chlormequat chloride on the proportion of phloem and xylem parenchyma in the storage root of carrot (Daucus carota L.)

A foliar application of gibberellic acid (GA) (100 mg/l) 5 weeks after emergence increased the shoot/storage root ratios, measured 4 weeks later, of 9 cultivars representing a range of types of carrot. Chlormequat chloride (CCC) (2000 mg/l) decreased the ratio for seven of the cultivars. Assessment 8 weeks after treatment showed fewer significant effects of both growth regulators.At the first harvest, GA decreased the phloem/xylem ratio and significantly reduced the amount of secondary phloem produced in the storage root of all cultivars. CCC had no comparable effect except for an increase in the phloem/xylem ratio for one cultivar at both harvests. The significance of these results is discussed in relation to secondary vascular tissue production in storage roots as affected by assimilate partitioning in the whole plant.     

Partitioning of 14C-Assimilate Between Sources and Sinks in the Monopodial Orchid Aranda Tay Swee Eng

Aranda Tay Swee Eng, an orchid hybrid with monopodial growthhabit, has a highly integrated source-sink assimilate partitioningpattern. The current inflorescence receives assimilate frommany rather than a few leaves, the vegetative apical shoot competeswith the inflorescence for assimilate supply in the upper shoot,and the fully-expanded leaves themselves constitute a majorsink for assimilate. Implications of these findings for improvementof harvestable yield in monopodial orchids are discussed. Aranda Tay Swee Eng, monopodial orchids, assimilate partitioning, source-sink relationships     

Root Confinement and its Effects on the Water Relations, Growth and Assimilate Partitioning of Tomato (Lycopersicon esculentum Mill)

Although it is well established that the root growth in manyspecies is very sensitive to mechanical impedance or to confinementin small volumes, little is known about the consequent effectson growth of the whole plant and the mechanisms involved. Thiswork investigated the effects of root confinement on the waterrelations, growth and assimilate partitioning of tomato (Lycopersiconesculentum Mill) grown in solution culture. Six-week old plants were transferred to either 4500 ml or 75ml containers filled with nutrient solution, and allowed togrow for 14 d. Transpiration, leaf-air temperature differences,and leaf diffusive resistances were measured frequently. Leaf,stem and shoot dry masses, leaf area and root length, were estimatedwhen the treatments were imposed and at the end of the experiment.After 14 d growth the root and shoot hydraulic resistances wereestimated from measurements of leaf water potential and transpirationrate, using a steady-state technique. Confining root growth to the small containers substantiallyreduced shoot and root growth and increased the proportion oftotal dry matter present in the stems. These effects were dueto drought stress. The hydraulic resistance of the root systemwas greatest in the confined plants. This led to more negativeleaf water potentials, increased leaf diffusive resistance,and reduced the net assimilation rate by a factor of 2.5. Transpirationper unit leaf area was less affected. However, cumulative transpirationwas also reduced by a factor of 2.5. mostly because of the smallerleaf area on the confined plants. Root hydraulic resistivitywas measured at 3.1 x 10¹²s m^–1 in the control treatment,but increased to 3.9 x 10¹² s m^–1 for roots in the smallcontainer. The mechanisms by which root confinement caused drought stressand disrupted the pattern of assimilate partitioning are discussedin detail. Assimilate partitioning, Lycopersicon esculentum, root confinement, plant growth, root growth, root resistance, shoot resistance, tomato, transpiration, water-use efficiency     

Sink development and dry matter distribution in storage root crops

Events during early development are examined for their contribution to dry matter distribution in the mature plant. It is concluded that timing of initiation of the storage root is not a major cause of differences between cultivars of carrot. These may airse from differences in the control of partitioning of assimilate at storage organ initiation. Root crop species may be affected differently by environmental factors such as density as a result of different thresholds of response of sink development to source stimuli.     

Dry Matter Distribution between the Shoot and Storage Root of Carrot (Daucus carota L.): I. Comparison of Varieties

The distribution of dry matter between shoot and storage rootof a wild variety and cultivars of carrot was investigated ina glasshouse experiment and a field experiment. In both experimentsthe deliberate introduction of variation in plant size revealeda linear relationship between the log weights of these partsat any time throughout growth. In the glasshouse experiment,in which growth was examined from approximately 10–600mg mean dry wt per plant, the slope of the relationship betweenIn shoot and In storage root dry wts decreased markedly duringdevelopment to 100 mg. Subsequently the slope became reasonablyconstant and the intercept progressively more negative. Datafrom the field experiment covered a similar timescale (48–115days) to the glasshouse experiment but the plants grew morequickly and were much heavier (180 mg–6 g dry wt). Therelationship between shoot and storage root for these data wassimilar to that for the plants of comparable size in the glasshouseexperiment. Over the range of development for which the slopeof the relationship was constant, data from both experimentswere described fairly well by the following equations: In shoot = – time+ In storage root where , and are constants. Between cultivars the slope ()and time constant () were similar, but the intercept () increasedwith maturity time of the cultivars. For wild carrot the slopewas larger and the time constant smaller than estimates forthe cultivars. These results on dry matter distribution arediscussed in relation to a model for partitioning assimilatein an individual plant. Daucus carota L., carrot, assimilate distribution, dry matter distribution, storage root, partitioning model     

Investigation of Quantitative Relationships in Dry Matter Distribution Between Tops and Storage Roots in Sugarbeet

Data from both controlled environment and field experimentson sugarbeet are used to support the concept that dry matterdistribution between top and storage root growth can be adequatelyquantified by over a large part of the growing season. Very young plants,prior to cambial ring development, had higher slopes for thelog-log relationship at each harvest. Estimates of the slopesbecame relatively constant about 30 d after emergence indicatingthat the pattern for dry matter distribution was determinedat an earlier stage of growth than is apparent from considerationof linear growth scales. Estimates of and are similar to thosereported for other storage root crops with differences in reflectingrelative size differences. Differences in applied N in the fieldwere detected as a shift in the intercept of the relationshipfor dry matter distribution. In general, the results with sugarbeetare agreement with the Barnes' model, which relates assimilatepartitioning to relative sink activities, respiration rates,and initial weights of plant parts. Beta vulgaris L., sugarbeet, dry matter distribution, assimilate partitioning, storage root, growth pattern, nitrogen, root/shoot weight     

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     

Growth and Development of Radish (Raphanus sativus, L.) Under Selected Light Environments

Plants of radish (Raphanus sativus L.) were grown under selectedlight conditions in controlled environmental chambers in orderto monitor the role of photoperiod, irradiance level and inputlight energy in plant development. Results indicated that thedaily input of light energy was the most important light factoraffecting leaf development while photoperiod and irradiancelevel had the major influences on storage organ development.Distribution of assimilates to leaves and storage organs variedunder different light regimes with long photoperiods and highirradiances producing the largest storage organs. Once initiated,the rate of storage organ growth was similar under all testedlight environments. Raphanus sativus L., radish, growth, development, light, photoperiod, assimilate distribution, storage organ