The Initiation and Growth of Axillary Bud Primordia in Relation to Flowering in Trifolium repens L.

The initiation and growth of axillary bud primordia in relationto the growth of their subtending leaves was observed at theapices of three clones (A. B. and C) of white clover grown invarious combinations of photoperiod and temperature. ClonesA, B, and C flower in response to low temperatures, and clonesA and C, but not B, in response to a transfer from short tolong photoperiods at higher temperatures. The rate of growth of buds and leaves from node to node waslittle influenced by the various treatments imposed, but theinitiation of axillary bud primordia relative to the apicaldome was stimulated in conditions conducive to flowering. The number of budless leaf primordia at the apex ranged froma maximum average of 2.25 at 20° C. to approximately o.8oat 10° C. in all three clones. At the higher temperatures,runners possessed 2.06 budless nodes in short days but only1.12 in long days in clones A and C. In clone B, daylength didnot influence bud initiation at the higher temperature. The results provide evidence of the homology between vegetativeand repro-ductive axillary bud primordia. It is suggested thatflowering is brought about by the removal of an inhibition withinthe apex which leads to the precocious initiation of axillarybud primordia. Following the initiation of axillary bud primordia, the resultsshow their growth to be uninhibited for 6-7 plastochrons. Rapidinflorescence development occurs during this phase. Apical dominancehas no apparent influence on vegetative axillary buds untilthe onset of rapid petiole elongation in their subtending leaves.     

Accumulation of myo -Inositol in Actinidia Seedlings Subjected to Salt Stress

Seedlings of kiwifruit (Actinidia deliciosa (A. Chev.) C. F.Liang et A. R. Ferguson vardeliciosa ) and A. arguta (Sieb.et Zucc.) Planch. ex Miq. grown in hydroponic nutrient solutionswith elevated salt (MgSO₄and KCl) concentrations showed visiblesigns of stress at salt concentrations of 50 m M and above.The polyol myo -inositol accumulated in leaf tissue when thesalt was added to 15 m M or more, with increases being similarin the two species. The increase in concentration of myo -inositolwas approximately linear with rising salt. At any given saltconcentration an increase in myo -inositol was linear with timefrom application of salt.myo -Inositol concentrations increasedwithin the first 24 h of salt treatment, and declined againas quickly once the stress was removed. Sucrose also increasedwith salt stress, accumulating only once plants showed physicalsigns of stress. Accumulation of myo -inositol was negativelycorrelated to fructose and glucose. Copyright 1999 Annals ofBotany Company Actinidia arguta, Actinidia deliciosa, kiwifruit, leaf tissue, myo -inositol, salt stress, sucrose.     

Changes in Cell Wall Composition and Water-soluble Polysaccharides During Kiwifruit Development

Changes in both cell wall and water-soluble polysaccharide compositionduring the growth of kiwifruits [Actinidia deliciosa (A. chev.) C. F. Liang and A. R. Ferguson var. deliciosa ‘Hayward’]were investigated. Cellulose was the major wall polysaccharide,with galactose and uronics the main non-cellulosic sugars. Muchsolubilization of cell wall pectic polysaccharides was detected.While wall-galactose solubilization started 3 months after anthesis,polyuronide degradation did not start until the fifth month,1 month prior to the harvest date. Parallel to these processes,a linear increase in water-soluble polysaccharides was detected.These mainly comprised galactose-rich polymers in the first3 months and little-branched polyuronides after the fifth month.Two different mechanisms for galactose and uronic acid solubilizationfrom kiwifruit cell walls during fruit development are proposed. Actinidia deliciosa ; cell wall; fruit growth; kiwifruit; water-soluble polysaccharides     

Winter bud content according to position in 3-year-old branching systems of 'Granny Smith' apple

An investigation was made of the number of preformed organs in winter buds of 3-year-old reiterated complexes of the 'Granny Smith' cultivar. Winter bud content was studied with respect to bud position: terminal buds were compared on both long shoots and spurs according to branching order and shoot age, while axillary buds were compared between three zones (distal, median and proximal) along 1-year-old annual shoots in order 1. The percentage of winter buds that differentiated into inflorescences was determined and the flowers in each bud were counted for each bud category. The other organ categories considered were scales and leaf primordia. The results confirmed that a certain number of organs must be initiated before floral differentiation occurred. The minimum limit was estimated at about 15 organs on average, including scales. Total number of lateral organs formed was shown to vary with both bud position and meristem age, increasing from newly formed meristems to 1- and 2-year-old meristems on different shoot types. These differences in bud organogenesis depending on bud position, were consistent with the morphogenetic gradients observed in apple tree architecture. Axillary buds did not contain more than 15 organs on average and this low organogenetic activity of the meristems was related to a low number of flowers per bud. In contrast, the other bud categories contained more than 15 differentiated organs on average and a trade-off was observed between leaf and flower primordia. The ratio between the number of leaf and flower primordia per bud varied with shoot type. When the terminal buds on long shoots and spurs were compared, those on long shoots showed more flowers and a higher ratio of leaf to flower primordia.     

Effect of light quality (red: far-red ratio) and defoliation treatments applied at a single phytomer on axillary bud outgrowth in Trifolium repens L.

We studied the effects of light quality and defoliation on the rate of phytomer appearance and axillary bud outgrowth in white clover. The treatments were applied to one phytomer, a phytomer being defined as the structural unit comprising a node, internode, axillary bud, subtending leaf and two nodal root primordia. Light of a low red:far-red (R:FR) ratio (0.27) was applied to a target phytomer either (i) within the apical bud and then to the axillary bud after emergence of the phytomer from the apical bud, or (ii) to the axillary bud only after emergence. The light conditions were directed to these specific parts of the plant by collimating light from small FR light-emitting diodes; with this technique we were able to change the light quality without any change in the level of photosynthetically active radiation. The subtending leaf of the target phytomer was retained or defoliated when it had emerged from the apical bud. FR light applied from the time the phytomer was within the apical bud caused a delay in branch appearance at the target phytomer. In contrast, direct treatment of the axillary bud with FR light after it had emerged from the apical bud did not result in any delay in branch appearance. As the light treatment of the apical bud may have changed the light environment of any of the organs contained in the bud we were unable to ascribe the delay in branch appearance to light perception by any particular organ. However, indirect evidence leads to the conclusion that the likely site of light perception was the developing leaf subtending the axillary bud while it was the outermost phytomer within the apical bud. These results do not support the hypothesis that the R:FR ratio of light incident at an axillary bud site is the environmental factor that controls bud development. Defoliation of the unfolding leaf reduced the rate of phytomer appearance on the main stolon but had no immediate effect on branch appearance. As a consequence there was a reduction in the number of phytomers between the stolon apical meristem and the first phytomer with a branch. This is frequently taken to indicate a relaxation of apical dominance, but in this case was found not to involve a direct effect on bud activity. A current model of white clover growth suggests that there is integration of activity between apical meristems but independence of activity and response to the local micro-environment by axillary buds. In contrast, we found that (i) defoliation reduced phytomer appearance only at the main stolon apical meristem and not at all the meristems in the plant and (ii) that a change in the local light environment of an axillary bud had no discernible effect on bud activity once the bud had emerged from the apical bud but could delay branching if applied before emergence. These results are at variance with the predictions of the model.     

Estimating the Bioenergetic Cost of a Developing Kiwifruit Berry and its Growth and Maintenance Respiration Components

A response surface was developed by regression analysis to quantifythe seasonal respiratory losses by a kiwifruit [Actinidia deliciosa(A. Chev.) C. F. Liang et A. R. Ferguson var. deliciosa cv.Hayward] berry growing in Fresno, CA. The equation of the surfacewas LNRESP = 1·622 + 0·0697 x TEMP –0·0472x DAY + 0·000165 x DAYSQ, where LNRESP is the naturallogarithm of the respiration rate (nmol CO₂ g d. wt^–1s^–1), TEMP is fruit temperature (°C), DAY is the numberof days after flowering, and DAYSQ is the square of the numberof days after flowering. Respiratory losses for a fruit witha final dry mass of 18·5 g were calculated to be 5·57and 5·92 g glucose per fruit per season in 1985 and 1986,respectively. Maintenance respiration was estimated to be 2·84and 3·19 g glucose per fruit per season for 1985 and1986, respectively. The total calculated bioenergetic cost ofkiwifruit berry growth and respiration was 25·25 and25·60 g glucose per fruit per season for 1985 and 1986,respectively. Respiratory losses, expressed as a proportionof the total carbohydrate required for fruit growth, were significant(mean 22·6%). The cost of fruit growth was estimatedto be very similar for two cooler sites (Davis and Watsonville)but estimates of maintenance respiration based on Fresno fruitrespiration data were unrealistically low for the Watsonvillesite. Actinidia deliciosa (A. Chev.) C. F. Liang et A. R. Ferguson var. deliciosa cv. Hayward, kiwifruit, growth respiration, maintenance respiration, bioenergetic costs, model     

Use of Interactive Computer Graphics for Simulation of Radiation Interception and Photosynthesis for Canopies of Kiwifruit Vines with Heterogeneous surface Shape and Leaf Area Distribution

A method incorporating interactive computer graphics to simulatespatially variable interception and canopy photosynthesis isdescribed. The method presents a graphical interface to a conventionalmodel of radiation interception and canopy photosynthesis. Includedis the capacity to consider a large number of positions withinthe canopy, thus providing a rapid and convenient representationof the dynamics of photosynthesis while also overcoming limitationsof one-dimensional models applied to complex plant canopies.The method was applied to examine spatial variability of photosynthesiswithin canopies of kiwifruit (Actinidia deliciosa) vines growingon two trellis types. The diurnal integral of simulated canopyphotosynthesis, assuming sunny conditions, for a vine trainedon a horizontal 'Pergola' trellis was 14% higher than that fora vine with similar leaf area distribution trained on a 'T-bar'trellis with inclined surfaces. Simulations of photosynthesisfor vines on a T-bar trellis, assuming spatially variable leafarea distributions as measured under filed conditions, indicateddisproportionate contributions from different regions of thecanopy. Canopy regions inclined to the east or the west wereusually the major sites for photosynthesis immediately aftersunrise and before sunset respectively, while regions near thecordon were the most important overall. For any day, the maximumsimulated photosynthetic rate generally declined with distancefrom the cordon and, at any distance from the cordon, increasedwith leaf index. For a vine with an average leaf area indexof 2·7, diurnal integrals of photosynthesis on a sunnyday in late summer ranged from 1·0 mol CO₂ m^-2 near thecordon to 0·5 mol CO₂ m^-2 at 1·5 m from the cordon.Within-canopy shading was more important on sunny days thanon cloudy days, while the spatial distribution of leaf areawas especially important on cloudy days. Comparison of simulationswith direct measurements of canopy photosynthesis indicatedthat a numerical integral of simulated photosynthesis, basedon a large number of canopy positions, provided a reasonableestimate of total canopy photosynthesis.Copyright 1993, 1999Academic Press Actinidia deliciosa, kiwifruit, interactive computer graphics, mathematical modelling, photosynthesis, radiation interception, spatial heterogeneity     

Growth and Compositional Changes in Kiwifruit Berries from Three Californian Locations

Kiwifruit [Actinidia deliciosa (A. Chev.) C. F. Liang et A.R. Ferguson var. deliciosa cv. Hayward] berries were sampledfrom three Californian locations, two sites in the central valley(near Davis and Fresno), the other on the coast (near Watsonville).Samples were analyzed for nitrogen, lipid, starch, soluble sugars,organic acids and ash, at regular intervals from flowering untilharvest. The results of the analyses of the fruit from the twocentral valley sites were similar, but markedly different fromthose from the cooler coastal site Sugar/acid ratios were consistentlyhigher in fruit from the coast than the central valley. Thismay prove a useful refinement to the current maturity index. Actinidia deliciosa (A. Chev.) C. F. Liang et A. R. Ferguson var. deliciosa cv. Haywood, kiwifruit, fruit growth, fruit composition, seasonal changes     

Effect of Light Quality (Red:Far-red Ratio) at the Apical Bud of the Main Stolon on Morphogenesis of Trifolium repens L.

A controlled environment experiment investigated whether thered:far-red (R:FR) ratio of light at the apical bud of the mainstolon could alter plant morphogenesis in clonal cuttings ofwhite clover (Trifolium repens L.) The apical bud included theapical meristem, five to six developing leaf primordia withassociated axillary bud primordia and stipules and the firstemerged folded leaf until development was greater than 0·3on the Carlson scale. Three light regimes were imposed on theapical bud by collimating light from R or FR light-emittingdiodes so that the R:FR ratio of light incident at the apicalbud was set at 0·25, 1·6 or 2·1, withoutsignificantly altering photosynthetically active radiation.The effect of these light regimes on white clover seedling growthwas also tested. At a low R:FR ratio seedling hypocotyl and cotyledon lengthswere significantly longer. However, with the cuttings, the lighttreatments did not alter node appearance rate or internode lengthof the main stolon, petiole length, area of leaves or totalshoot dry matter. There was one significant photomorphogeneticresponse in the cuttings, a delay of 0·5 of a phyllochronin the appearance of branches from axillary buds in the lowR:FR ratio treatment relative to the other treatments. Wherebranch appearance was delayed plants had fewer branches. Thisdifference could be ascribed solely to a delay in branch appearanceas there were no significant treatment effects on either theinitiation of axillary bud primordia within the apical bud,the probability of branching or on the rate of growth of branchesafter appearance. Because treatment of the apical bud inducedonly one of the many previously observed responses of whiteclover to a decrease in the R:FR ratio of light, we concludethat other plant organs must also sense the quality of incidentlight.Copyright 1994, 1999 Academic Press White clover, Trifolium repens, apical bud, light quality, red:far-red ratio, light-emitting diode, branching, axillary buds, photomorphogenesis     

Localization of axillary meristems during different stages of radish ontogenesis

An analysis of axillary meristem (axillary bud) localization of radish (Raphanus sativus L. cv. Tetra-I?ówiecka) was undertaken on vernalized (flowering) and unvernalized (vegetative) plants. It has been shown that the localization of these meristems can be different on successive nodes of the same plant and is connected with the development stages of the plants. The axillary meristems can arise on the stem as well as in the leaf axil or on the base of the subtending leaf. The localization of axillary meristems has been discussed in relation to growth directions and growth correlations inside the meristematic region of the shoot apex.     

Comparison of Four Methods Calculating the Seasonal Pattern of Plant Growth Efficiency of a Kiwifruit Berry

Four methods of determining the substrate requirements for synthesisof a kiwifruit [Actinidia deliciosa (A. Chev.) C. F. Liang etA. R. Ferguson var. deliciosa cv. Hayward] berry were comparedusing data derived from common kiwifruit berry samples collectedfrom anthesis to fruit maturity. The four methods were basedon fruit proximal analysis, elemental analysis, heats of combustion,or tissue carbon content. All methods gave similar patternsof seasonal costs and values of final cost to the plant (mean1.21 g glucose g^–1 season^–1) but there was lessagreement for growth respiration (mean 0.147 g glucose g^–1season^–1). This is the first time that a continuous recordof growth cost over the course of development has been presented,and the trends in seasonal cost reflect the uptake into andsynthesis of the different biochemical constituents in the fruit.The differences between the results of each method reflect theunderlying assumptions used in their development. It appearsfrom this work that the method of McDermitt and Loomis (1981),utilizing elemental analyses, is most preferred. Actinidia deliciosa (A. Chev.) C. F. Liang et A. R. Ferguson var deliciosa cv Haywood, kiwifruit, true growth yield, plant growth efficiency, production value, glucose value, bioenergetic cost     

Regulation of Branching in Decussate Species with Unequal Lateral Buds

In the decussate plants Alternanthera philoxeroides and Hygrophilasp. the opposite axillary bud primordia are of unequal sizefrom the time of their inception; the larger or + buds lie alongone helix and the smaller or – buds along another (helicoidalsystem). In decapitated plants of Alternanthera both buds grewout, but unequally; if the node was vertically split growthof the two shoots was more equal, and if the + buds were excisedgrowth of the – shoots approximately equalled that ofcontrol + shoots. In decapitated shoots of Hygrophila grownin sterile culture only one bud, the + or larger one, grew outat each of the upper nodes. In excised cultured nodes, also,only the + bud grew out; but if the nodes were split longitudinallyboth buds grew out, initially rather unequally. These experimentssupport the view that the regulation of branching in these specieshas two components, apical dominance and the dominance of thelarger (+) bud over the smaller (–) bud at the same node.The restriction of growth potentiality imposed on the –bud is not permanent but can be modified. Further correlativeeffects on bud outgrowth include those of the subtending leavesand of buds at other nodes.     

Determination for inflorescence development is a stable state,separable from determination for flower development in Pisum sativum L. buds

Terminal meristems of Pisum sativum (garden pea) transit from vegetative to inflorescence development, and begin producing floral axillary meristems. Determination for inflorescence development was assessed by culturing excised buds and meristems. The first node of floral initiation (NFI) for bud expiants developing in culture and for adventitious shoots forming on cultured meristems was compared with the NFI of intact control buds. When terminal buds having eight leaf primordia were excised from plants of different ages (i.e., number of unfolded leaves) and cultured on 6-benzylaminopurine and kinetin-supplemented medium, the NFI was a function of the age of the source plant. By age 3, all terminal buds were determined for inflorescence development. Determination occurred at least eight nodes before the first axillary flower was initiated. Thus, the axillary meristems contributing to the inflorescence had not formed at the time the bud was explanted. Similar results were obtained for cultured axillary buds. In addition, meristems excised without leaf primordia from axillary buds three nodes above the cotyledons of age-3 plants gave rise to adventitious buds with an NFI of 8.3 ±0.3 nodes. In contrast seed-derived plants had an NFI of 16.5 ±0.2. Thus cells within the meristem were determined for inflorescence development. These findings indicate that determination for inflorescence development in P. sativum is a stable developmental state, separable from determination for flower development, and occurring prior to initiation of the inflorescence at the level of meristems.     

Vibratory Collection of Actinidia deliciosa (Kiwifruit) Pollen

Dry, powdery pollen grains were expected from 'buzz pollinated'flower species. However, vibration of Actinidia deliciosa (kiwifruit)anthers (a buzz pollinated species) by a mechanical shaker,at similar vibrations to Bombus terrestris caused clumps ofpollen joined by small droplets of tapetal fluid to be ejected.Pollen that was largely covered with tapetal fluid could notbe removed by vibration, whereas dehydrated pollen was easilyremoved, even without vibration. The late desiccation of A.deliciosa anthers after anthesis meant that pollen removed byvibration depended on the anther maturity. The presence of thedroplets gives insects which vibrate while foraging advantagesover insects which do not. A vibration attachment on a commercialpollen harvester increased the weight of pollen collected by57% over the whole day; 91% in the afternoon.Copyright 1994,1999 Academic Press Actinidia deliciosa, Bombus spp., bumblebees, buzz pollinate, kiwifruit, pollen collection, tapetal fluid, vibration     

Sensitivity of Floral Shoot Growth, Fruit Set and Early Fruit Size inActinidia deliciosato Local Carbon Supply

The sensitivity of pre-anthesis shoot growth, fruit set andearly fruit size in kiwifruit [Actinidia deliciosa(A. Chev.)C. F. Liang et A. R. Ferguson] to cane girdling and defoliationtreatments was tested, and the results discussed in terms ofshoot carbon balance. The nature and degree of responses obtainedwas dependent on the time of application. When treatments wereapplied soon after budbreak, defoliation initially reduced growth,whereas increasingly severe girdling treatments induced responsesprogressing from stimulation to severe impairment of growthand photosynthetic capability. Similar treatments imposed laterin shoot development produced qualitatively different results.The development of floral tissues on the shoot is shown to besensitive to shoot carbon status during a period close to anthesis,and the significance of this result in relation to fruit heterogeneityat maturity is discussed.Copyright 1998 Annals of Botany Company Actinidia deliciosa, kiwifruit, shoot growth, fruit variability, fruit set, reserves.     

Effects of Water Stress on Fruit Quality Attributes of Kiwifruit

Four-year-old kiwifruit vines (Actinidia deliciosa(A. Chev.)C. F. Liang et A. R. Ferguson var.deliciosacv. Hayward) werestudied to determine response of the plant and effects on fruitquality when irrigation water was withheld either early or latein the growing season. The greatest effect on fruit growth occurredwhen water was withheld early in the season. Harvest weightof fruit from early-stressed vines was approx. 25% less thanthe weight of fruit on control vines. Early season water stressresulted in a transient increase in concentrations of solublecarbohydrates in both leaves and fruit. This was accompaniedby a reduction in stomatal conductance of the leaves. Starchlevels in leaves but not fruit were reduced by both stress treatments.Concentrations of sucrose at harvest in fruit from vines stressedlate in the season were markedly higher than in other fruit,and softness of the fruit was unaffected. These differenceswere maintained through the 12 weeks in cool storage after harvest.Withholding irrigation water to kiwifruit vines late in theseason may prove a useful management tool to manipulate somequality attributes of the fruit.Copyright 1998 Annals of BotanyCompany Kiwifruit;Actinidia deliciosa; water stress; fruit quality; soluble solids.     

Initiation of axillary and floral meristems in Arabidopsis

Shoot development is reiterative: shoot apical meristems (SAMs) give rise to branches made of repeating leaf and stem units with new SAMs in turn formed in the axils of the leaves. Thus, new axes of growth are established on preexisting axes. Here we describe the formation of axillary meristems and floral meristems in Arabidopsis by monitoring the expression of the SHOOT MERISTEMLESS and AINTEGUMENTA genes. Expression of these genes is associated with SAMs and organ primordia, respectively. Four stages of axillary meristem development and previously undefined substages of floral meristem development are described. We find parallels between the development of axillary meristems and the development of floral meristems. Although Arabidopsis flowers develop in the apparent absence of a subtending leaf, the expression patterns of AINTEGUMENTA and SHOOT MERISTEMLESS RNAs during flower development suggest the presence of a highly reduced, "cryptic" leaf subtending the flower in Arabidopsis. We hypothesize that the STM-negative region that develops on the flanks of the inflorescence meristem is a bract primordium and that the floral meristem proper develops in the "axil" of this bract primordium. The bract primordium, although initially specified, becomes repressed in its growth.     

The induction of sun and shade leaves of the European beech (Fagus sylvatica L.): anatomical studies

Summary Primordia from buds of sun and shade twigs of European beech (Fagus sylvatica L.) were collected six times a year for anatomical investigations. Differentiation into sun-leaf and shade-leaf primordia was first observed in early August. Sun-leaf primordia had five, and shade-leaf primordia four layers of mesophyll meristem cells. With potted graft unions of beeches possible structural changes of leaf primordia were investigated. Trees adapted to shade develop sun-leaf primordia when put into full daylight, provided the transfer happened before July. Trees adapted to full daylight developed leaf primordia which remained structurally sun-leaf primordia when the plant was kept under shade conditions. Shadeleaf branches of young beech trees cut in February in order to expose the shade buds to full daylight developed either shade leaves or intermediate shade/sun leaves. These experiments show that the subtending leaf may provide the developing axillary bud with photoassimilates, but its character, whether sun or shade leaf, has no influence on the character of the developing leaf primordia.     

A Method for Analysing Plant Architecture as it Relates to Fruit Quality Using Three-dimensional Computer Graphics

A method was developed for the spatial analysis of plant architectureas it relates to the within-plant variation in the physical,chemical, and postharvest characteristics of the fruit Computergraphics were used to reconstruct the architectural frameworkand spatial arrangement of the fruit in the canopy of kiwifruitvines (Actinidia deliciosa) trained on two different supportstructures An infra-red beam theodolite was used to obtain thespatial coordinates of the vines components The data files generatedby the theodolite were in turn used with software specificallywritten for the project (MAPIT—Microcomputer Aided PlantImaging Technology) to provide a 3-dimensional reconstructionof the original vines Each fruit was colour coded so that extremesin their attributes could be easily identified and accuratelylocated in the canopy of the vine Patterns were clearly discerniblefor both the pergola and T-bar trained vines The heavier fruitwere located at the apical ends of the canes, while greatersoluble solids concentrations were associated with the smallerfruit located closer to the cordon These patterns were consistentfor all of the vines examined The use of the theodolite coupledwith the computer graphics described in this paper providesa rapid and objective means of accurately describing plant architecture Computer graphics, plant architecture, spatial analysis, theodolite, three-dimensional analysis, fruit quality, Actinidia deliciosa, kiwifruit