Proline, Hydroxyproline, and Lily Pollen Tube Elongation

Cytoplasm of freshly-harvested, ungerminated Lilium longiflorum,cv. Ace pollen contains 0.14 per cent soluble and 0.35 per centprotein-bound proline (pro). Their metabolic fates in germinationand tube elongation are not known with certainty. Here is reportedconversion of pro to hydroxyproline (hyp)—containing constituentsas well as distribution and isolation of these constituents.Colorimetry revealed pro and hyp in wall, trichloroacetic acid(TCA)—precipitable, and TCA-soluble cytoplasmic fractions.A balance sheet summarizing quantitative changes in pro andhyp for these fractions revealed that TCA—precipitablecytoplasmic pro could be a precursor to wall-bound pro and asubstrate for hydroxylation yielding cytoplasmic and wall-boundhyp. To determine whether hyp was a component of tube and/orgrain walls, pollen was allowed to germinate 1.5 h and thentransferred to sorbitol medium which prevented further tubeelongation. Hyp was absent from walls of transferred pollen.Electron microscope autoradiography of tubes exposed to ²H-prosuggested that a pro- and/or hyp-containing constituent waslocalized in the growing tip. Light microscope autoradiographyof intact tubes labelled with ¹⁴C-pro showed that the constituentwas distributed throughout the pollen tubes. Gel filtrationof hyp-containing material enzymically released from walls supportedthe view that they contained hyp-glycopeptides.     

The Structure and Function of Pine Forest in Central Himalaya. I. Dry Matter Dynamics

The present study deals with structure and function of fourareas of Himalayan chir pine forest. Tree layer was monospecificon all sites with varied density and basal cover in the rangeof 540–1630 individuals per ha and 25·0–47·2m²ha^–1, respectively. Shrubs having low density were sparselydistributed. All allometric equations relating to biomass ofdifferent components, to circumference at breast height (cbh)were significant, with the exception of that for cone biomass.Total vegetation biomass (115–236 t ha^–1) was distributedas 113–283 t ha^–1 in trees. 0·56–0·82t ha^–1 in shrubs and 1·63–2·57 t ha^–1in herbs. Total forest floor biomass including herbaceous litterranged between 9·6 and 13·6 t ha^–1. Of thetotal annual litter fall (4·26–7·38 t ha^–1),60·3–75·1 per cent was distributed in leaflitter and 24·9–39·7 per cent in wood litter.Turnover rate of tree litter varied from 0·45 to 0·53,whereas rates for shrubs and herbs were assumed to 1. Net primaryproduction of total vegetation ranged between 9·91 and21·2 t ha^–1 year^–1, of which the contributionof trees, shrubs and herbs was 76·5– 88·1per cent 0·6–1·8 per cent and 11·3–21·5per cent, respectively. A compartment model of dry matter onthe basis of mean data across sites was developed to show drymatter storage and flow of dry matter within the ecosystem. Pinus roxburghii forest, biomass, litter fall, net primary production, compartmental transfer     

Post-transcriptional Control of Nitrate Reductase Formation in Green Algae

Cycloheximide (2·0 µg ml^–1) inhibits theincorporation of [¹⁴C]phenylalanine and [¹⁴C]adenine into insolublecompounds in Ankistrodesmus braunii. 6-Methylpurine (1·0mM) inhibits only the incorporation of [14C]adenine and it isconcluded that it inhibits RNA synthesis. When ammonium-growncells of Ankistrodesmus or Chlorella are nitrogen-starved orwhen ammonium-grown cells of Dunalitlla are resuspended in nitratemedium, the appearance of nitrate reductase in these organismsis not inhibited by 6-methylpurine. The appearance of nitratereductase activity in Ankistrodesmus or Chlorella is inhibitedby 6-methylpurine when ammonium-grown organisms are preincubatedwith this substance for 1-2 h before nitrogen starvation. Itis concluded that cells growing with ammonium and lacking nitratereductase activity nevertheless contain preformed mRNA for nitratereductase synthesis.     

Distribution of 14C Assimilates in the Cowpea (Vigna unguiculata L.) in Relation to Fruit Abscission and Treatment with Benzyladenine

The distribution of ¹⁴C assimilates from ¹⁴C-sucrose was studiedin relation to premature fruit abscission in two cowpea cultivars,Adzuki and Mala. In both cultivars most of the radioactivitywas recovered in the fruits, constituting 63–85 per centof the total ¹⁴C imported from the fed leaflet. This was followedby the root, leaves and stem in descending order, except thatin Mala, import by the stem was greater than that by the leaves.Adzuki imported 56 per cent more ¹⁴C than Mala, from the fedleaflet. In Adzuki, which exhibits a relatively low degree ofabscission of young fruits, the ratio of ¹⁴C accumulated bypeduncle 1 (oldest) fruits to that of peduncle 3 (youngest)fruits was 0·31; while in Mala it was 0·61. Ratiosof the combined accumulation by peduncles 1 and 2 fruits topeduncle 3 fruits were 0·81 for Adzuki, and 1·88for Mala. The more mature fruits of Mala thus constituted amore potent sink for ¹⁴C assimilates than those of Adzuki. In Adzuki, benzyladenine treatment of young fruits at each pedunclewas not significantly effective in reversing or modifying thenormal gradient of assimilates in fruits of different ontogeny.However, in Mala, BA treatment of the youngest fruits caused43 per cent increase in ¹⁴C import, when compared with correspondingfruits of control plants. In Adzuki, BA had no significant effecton total fruit weight, whereas in Mala the weight was increasedby about 36 per cent.     

Investigations of Carbon Transport in Plants: II. THE EFFECTS OF LIGHT AND DARKNESS AND SINK ACTIVITY ON TRANSLOCATION

The use of ¹¹C as a tracer has allowed repetitive measurementsof the speed of assimilate translocation to be made on singlemaize plants throughout prolonged periods of light and darkness.The speed appeared to double when the light was switched on.The time required to achieve a maximum speed, usually about3·5 cm min^–1, depended on the duration of the previousdark period. When the plant was transferred to darkness thespeed immediately decreased by about 20 per cent and continuedto decrease over the next 20 h to values of 0·5 to 0·9cm min^–1. The mean speed of translocation in tomato in the light, andother C3 plants, was usually about 1 cm min^–1. It wasreduced by 15–30 per cent when the fruit was removed orcooled from 26 to 10°C.     

The Carbon Economy of Rubus chamaemorus L. II. Respiration

Respiratory activity and seasonal changes in carbohydrate contentof the storage organs of Rubus chamaemorus L. have been investigated.Leaf dark respiration rate increases in a non-linear mannerfrom 0·7 mg CO₂ evolved dm^–2 h^–1 at 0 °Cto 4·6 rng CO₂ evolved dm^–2 hh^–1 at 30 °C.Root and rhizome respiration rates increase from 1 µ1O₂ uptake g^–1 fresh weight h^–1 at 0.7 ° C to10 µ10, uptake g^–1 f. wt h^–1 at 20 °C.Rhizome carbohydrate reserves decline from a September peakof 33 per cent alcohol insoluble d. wt to 16 per cent in May. The circumpolar distribution of R. chamaemorus is discussedin relation to the evidence presented here and in the precedingpaper of the series.     

The Transport of Sugars in Developing Fruits of Satsuma Mandarin

Transport of sugars to the juice sacs of developing satsumamandarin (Citrus unshui Marc) has been studied in attached fruitsand in isolated fruit pieces. ¹⁴CO₂ fed to the leaves resultedin [¹⁴C]sugar accumulation in the juice sacs, mainly as [¹⁴C]sucrose.Uptake of sucrose and glucose by the excised fruit pieces proceededlinearly with time. Sucrose uptake was linearly related to sucroseconcentration over the range 25–300 mM, with no indicationof saturation. This uptake was insensitive to pH (5, 7 or 9),Ca²⁺(3 mM), PCMBS (2.5 mM), DNP (1 mM) or vanadate (0.1 mM)but was slightly reduced by erythrosin (21 % by 0–1 mM;27 % by 1 mM). No competitive effect of glucose (up to 100 mM)was detectable on sucrose uptake from 100 mM solution. Mostof the [¹⁴C]sucrose uptake observed was reversible, althoughconsiderable hydrolysis and metabolic conversion were evidenced.A vanadate-sensitive ATPase was demonstrated by EM localizationon the plasma membrane of the juice sac cells. These resultsare interpreted in relation to the accumulation of assimilatesby the developing fruit. Transport: sugar, satsuma mandarin, juice sacs     

Germination of Sporangia of Phytophthora palmivora (Butl.) Butl.

Sporangia of Phytophthora palmivora germinated by either forminggerm tubes or producing zoospores. Two distinct modes of germ-tubedevelopment have been described. Sporangia in distilled waterformed zoospores at 10–34°C with an optimum at 22°Cbut germinated by means of germ tubes at 30 and 34°C only.Zoospore formation was inhibited to varying degrees by cocoapod extract, I.0 per cent (w/v) peptone and yeast extract, 100–500µg1^-1 thiamine, and by very low concentrations of severalamino acids, carbohydrates, and inorganic salts. Germ-tube formation was encouraged at 22°C by 1'0 per cent(w/v) peptone and yeast extract, by cocoa pod extract and exudate,10mM CaCl2, 1–10 mM MgSO₄. 7H₂O, 0.5 per cent (w/v) fructose,galactose, glucose, lactose, maltose, and sucrose, by 100 ppmarginine, aspartic acid, glutarnic acid, glycine, leucine, andtryptophane, and by 100–500 µg 1^-1 thiamine.     

Cytokinin-Mediated Translational Control of Protein Synthesis in Cultured Cells of Glycine max

Polyribosome formation was stimulated by cytokinin treatmentof cultured cells of Glycine max cv. Funk Delicious. When suspensioncultures were given 0·5 µM zeatin after 24 h inculture in medium lacking a cytokinin, a nearly 2-fold increasein the polyribosome/monoribosome ratio occurred over the subsequent3 h. The effect of actinomycin D and of 5-fluorouridine on RNAsynthesis and on the polyribosome/monoribosome ratios of thesecells was examined. Actinomycin D at 5 and 20 µg/ml^–1inhibitedtotal RNA synthesis by 39 and 60%, respectively, as measuredby [³H]uridine incorporation into acid-precipitable material.The degree of inhibition of precursor incorporation into polyribosomalRNA was similar. At 0·1 mM, 5-fluorouridine inhibited[³H]uridine incorporation by 76%, and [³H]guanosine incorporationby 66% into polyribosomal RNA after 3 h of treatment. Fractionationof the polyribosomal RNA by oligo(dT)-cellulose chromatographydemonstrated that low concentrations of both actinomycin D (5µg ml^–1) and 5-fluorouridine (0·1 mM) inhibitedthe synthesis of ribosomal RNA to a greater extent than thepoly(A)-containing fraction of the messenger RNA. Synthesisof the poly(A)-containing RNA was inhibited by 24% with 5µgml^–1 actinomycin D and by 30% with 0·1 mM 5-fluorouridine.At the above concentrations, these two inhibitors reduced thepolyribosome/monoribosome ratio of the cytokinin-deprived cellsover a 3 h period, but they did not prevent cytokinin-inducedpolyribosome formation. These results provide further evidencethat cytokinin regulates polyribosome levels through an effecton protein synthesis at the translational level     

Studies on the Growth in Culture of Plant Cells: XV. UPTAKE AND UTILIZATION OF URIDINE DURING THE GROWTH OF ACER PSEUDOPLATANUS L. CELLS IN SUSPENSION CULTURE

[2-¹⁴C]-uridine is rapidly taken up by sycamore cells in suspensionculture. A proportion of the radioactivity enters RNA withoutmeasurable delay, whilst the remainder equilibrates with a largepool of phosphorylated compounds, the major radioactive componentof which is 5'-UMP. Both the uracil and cytosine residues ofRNA receive label from [¹⁴C]-uridine and, when the cells aresupplied with high concentrations of uridine, these bases arederived almost exclusively from the nucleoside. [¹⁴C]-uridine is incorporated into RNA at all stages of thegrowth cycle of batch cultures; its continuing incorporation,when the total RNA content of the cells is rapidly decreasing,indicates a high rate of turnover of the total RNA. Long-termlabelling experiments also indicate turnover of RNA during thephase of active cell division and suggest that a large proportionof the degradation products are not re-utilized for RNA synthesis. Sycamore cells degrade [2-¹⁴C]-uridine with release of ¹⁴CO₂.The proportion degraded increases from 25 per cent at an externaluridine concentration of 10^–6M to 75 per cent at 10^–3M. Despite this, nucleic acids are the only macromolecules thatreceive a significant amount of radioactivity from [2-¹⁴]C-uridine.     

Effect of {alpha}-Tomatine on the Response of Wheat Coleoptile Segments to Exogenous Indole-3-acetic Acid

A concentration of 10^–5 M tomatine had no effect on leakagefrom, or elongation of, wheat coleoptile segments, but consistentlyreduced IAA-enhanced extension growth by c. 50 per cent. Therewas no evidence of chemical interaction between the alkaloidand the auxin in solution, and IAA action was not affected bypre-treatment for up to 3 h with 10^–5 M tomatine. Studieswith [2-¹⁴C]IAA revealed that 10^–5 M tomatine did notinhibit uptake of auxin into segments. The effect of pre-treatingsegments for up to 3 h with IAA could be virtually nullifiedby 10^–5 M tomatine, as could also IAA-induced changesin properties of coleoptile cell walls. Results are discussedin relation to the ability of tomatine to disrupt membrane functionand to current hypotheses implicating membranes in the primaryaction of auxin.     

Kinetics of Integration of Xyloglucan into the Walls of Suspension-Cultured Rose Cells

To study the kinetics of synthesis, wall-binding and degradationof xyloglucan, we incubated suspension-cultured rose cells for0–5–24 h in L-[1-³H]arabinose. >95% of the [³H]arabinosewas taken up within 2 h. UDP-Pentoses were maximally labelledwithin 0–5 h and had lost most of their ³H by 2 h afterthe addition of [³H]arabinose. Therefore, the 24 h experimentresembled a pulse-chase rgime. The [³H]xyloglucan formed wasfractionated into four cellular pools [detergent-extractable(interpreted as cytoplasmic), and guanidinium thiocyanate-,06 M NaOH- and 60 M NaOH-extractable (interpreted as progressivelymore firmly wall-bound)]; soluble extracellular xyloglucan wascollected as a fifth pool. All five pools of xyloglucan hadstarted accumulating ³H at their respective maximal rates by     

Metabolism of Pyrimidines in Protoplasts from Cultured Catharanthus roseus Cells

The metabolism of [2-¹⁴C]thymine, [2-¹⁴C]thymidine, [2-¹⁴C]uraciland [¹⁴C]uridine was investigated in protoplasts obtained fromsuspension cultures of Catharanthus roseus. Most of the exogenouslysupplied thymine, thymidine and uracil was degraded, and salvageof these pyrimidines accounted for 5–36 per cent of thetotal amount of ¹⁴C-labelled precursors which was metabolized.However, more than 80 per cent of the labelled uridine was utilizedfor the biosynthesis of nucleotides and nucleic acids, and therest was degraded. In contrast to the results from protoplastsof sugar cane cells in suspension culture, the data indicatethat protoplasts possess a pathway for the degradation of pyrimidines,and that the overall metabolism of these pyrimidines in protoplastsis very similar to the metabolism in the intact cells. Catharanthus roseus, madagascar periwinkle, protoplasts, pyrimidine metabolism     

Radiolabelling Studies on the Lipid Metabolism in the Marine Brown Alga Dictyopteris membranacea

The lipid metabolism of the marine brown alga D. membranaceawas investigated using [2–¹⁴C]acetate, [1–¹⁴C]myristate,[l–^I4C]oleate and [l–¹⁴C]arachidonate as precursors.On incubation with [2–¹⁴C]acetate, 18:1 and 16:0 werethe main products formed by de novo synthesis and incorporatedinto polar lipids. With all the exogenous substrates used, DGTAwas strongly labelled and the subsequent rapid turnover of radioactivitysuggested a key role for this lipid in the redistribution ofacyl chains and most likely also in the biosynthesis of theeukaryotic galacto-lipids produced in the absence of PC. Inthe glycolipids a continuous accumulation of radioactivity wasobserved with all the substrates used. The labelling kineticsof molecular species of MGDG suggested the desaturation of 18:1to 18:4 and of 20:4 (n-6) to 20:5 (n–3) acids on thislipid. Both PG and PE were primary acceptors of de novo synthesizedfatty acids and exogenous [l–¹⁴C]oleate, but no evidenceexists for a further processing of acyl chains on these lipids.TAG, although strongly labelled with all exogenous [l–¹⁴CJacids,was not labelled when [2–¹⁴C]acetate was used as a precursorindicating the flux of endogenous fatty acids to be differentof that of exogenously supplied fatty acids. (Received November 4, 1997; Accepted February 23, 1998)     

Radiolabelling Studies of Lipids in the Marine Cryptomonad Chroomonas salina in Relation to Fatty Acid Desaturation

Cells of Chroomonas salina were exposed to [¹⁴C]acetate, [^l4C]16:0,[¹⁴C]18:0, [¹⁴C]18:1(n-9), [¹⁴C]18:2(n–6) or [¹⁴C]18:3(n–3)for 1 h and then incubated for 24 h in non-radioactive medium.At the end of the pulse period, non-glycolipid polar lipidscontained the highest proportions of radioactivity incorporatedfrom [¹⁴C]acetate and [¹⁴C]18:3(n–3) whereas with [¹⁴C]16:0,[¹⁴C]18:1 and [¹⁴C]18:2(n–6), triacylglycerols were mosthighly labelled. ¹⁴C-18:0 was recovered mainly as non-esterifiedfatty acid. Monogalactosyldiacylglycerol initially contained17% of label incorporated from [¹⁴C]acetate but less than 3%of that from [¹⁴C]fatty acids. With all substrates, excluding[¹⁴C]18:0, a gradual transfer of label from polar lipids totriacylglycerols was observed during the chase period. Saturatesand monoenes synthesised from [¹⁴C]acetate were mostly transferedfrom phospholipids and glycolipids to neutral lipid withoutfurther desaturation. Most of the incorporated ¹⁴C-fatty acidsremained unchanged and only with [¹⁴C]18:3(n–3) was substantialamounts of label recovered in penta- and hexaenoic fatty acids.The results indicate that, under the conditions of the study,lipid synthesis in the algae was heavily dominated by triacylglycerolformation and that the mechanisms of fatty acid desaturationin this species may differ from those in higher plants. (Received December 10, 1991; Accepted March 6, 1992)     

Flavanoids in Pollen and Stigma of Brassica oleracea and Their Effects on Pollen Germination In Vitro

Brassica oleracea pollen was applied to a basic medium of 1.5per cent agar and 15 per cent sucrose to which flavanoids wereadded at three concentrations. Two types of agar were used;with agar 1, quercetin at a concentration of 0.5 x 10^–3per cent gave an increase in percentage germinating grains.With agar 2, an increase in germination occurred with kaempferoland naringin at concentrations of 0.5 x 10^–3 and 0.5 x10^–1 per cent respectively. Increase in pollen tube lengthoccurred with agar 2 and quercetin at a concentration of 0.5x 10^–3 per cent. The stigma tissue of B. oleracea contains at least three andthe pollen at least one glycoside of quercetin. The sugars inthe glycosides were not identified. Pollen germination and pollentube extension were not stimulated exclusively by the flavanoidspresent in the stigma. The flavanoid composition of the stigmadid not vary amongst five different S-allele genotypes, indicatingthat flavanoids are probably not directly involved in the incompatibilityreaction of B. oleracea.     

The Effect of Cycloheximide on Absorption of Ions by Bean (Phaseolus vulgaris L) Roots

10^–7 M cycloheximide inhibited bean (Phaseolus vulgaris)root elongation by about 20 per cent but it inhibited absorptionof rubidium, sodium, and phosphate ions to a much greater extent(34–71 per cent). Tips of intact plant roots grown inthe inhibitor showed more inhibition in ion uptake than adjacentproximal portions of the same roots and this is taken to indicatethat 10^–7 M cycloheximide does not exert its effect onion uptake by any uncoupling action. Sodium uptake from 0.5 or 10 mM NaCl solutions by root tipswas inhibited by 10^–7 M cycloheximide to twice the extentthat it was in the elongating region of the root. Assuming thatthe inhibitor affects the plasmalemma more than the tonoplast,Epstein's model of parallel operation of system 1 and system2 at the plasmalemma is supported.     

Carbon Translocation in the Tomato: Pathways of Carbon Metabolism in the Fruit-

The rate of carbon import by tomato fruits has been relatedto their carbon metabolism by examining the effects of fruittemperature on the metabolism of imported assimilates. ¹⁴C–sucrose,–glucose, –fructose, –malic acid and –citricacid were injected individually into young growing tomato fruitswhich were subsequently maintained at 25 or 5 °C for 48h. Fruit temperature greatly affected the proportions of ¹⁴Clost from the fruits by export and respiration. Only 40 percent of the injected ¹⁴C from ¹⁴C–sugars and 20 per centfrom ¹⁴C–acids was recovered from fruits at 25 °C.Less than 10 per cent of the injected ¹⁴C was exported, thebalance being respired. In contrast, more than 50 per cent ofthe injected ¹⁴C was recovered from cooled fruits, in whichthe import rate of carbon was presumably reduced, and 20–36per cent of injected ¹⁴C was exported. Cooling enhanced thesynthesis of ¹⁴C–sucrose from injected ¹⁴C–hexosesand inhibited the incorporation of ¹⁴C into starch and insolubleresidue. When ¹⁴C–sugars were injected, radioactivityexported from the cooled fruits was detected as sucrose in thephloem of the peduncles; radioactivity was also detected instems and roots when fruits were cooled. In almost fully–grownfruits injected ¹⁴C–compounds were metabolized less readilythan in smaller fruits. Conversion of ¹⁴C–hexoses to ¹⁴C–sucrosewas again enhanced by cooling (5 °C, but was less in fruitsmaintained at 35 °C than in controls. Lycopersicon esculentum, tomato, fruit, translocation, carbon metabolism     

Effect of Temperature on Photosynthesis and Photorespiration of Wheat Leaves

Wheat plants were grown in a controlled environment with daytemperatures of 18 ?C and with 500 µ Einsteins m^–28^–1 of photosynthetically active radiation for 16 h. Beforeanthesis and 2 to 3 weeks after, rates of net photosynthesiswere measured for leaves in 2 or 21% O₂ containing 350 vpm CO₂at 13, 18, 23, and 28 ?C and with 500 µEinsteins m^–2s^–1 of photosynthetically active radiation. Also, underthe same conditions of light intensity and temperature, therates of efflux of CO₂ into CO₂-free air were measured and,for mature flag leaves 3 to 4 weeks after anthesis, gross andnet photosynthesis from air containing 320 vpm ¹⁴CO₂ of specificactivity 39?7 nCi µmol^–1. When the O₂ concentration was decreased from 21 to 2% (v/v)the rate of net photosynthesis increased by 32 per cent at thelowest temperature and 54 per cent at the highest temperature.Efflux of CO₂ into CO₂-free air ranged from 38 per cent of netphotosynthesis at 13 ?C to 86 per cent at 28 ?C. Gross photosynthesis,measured by the ¹⁴C assimilated during 40 s, was greater thannet photosynthesis by some 10 per cent at 13 ?C and 17 per centat 28 ?C. These data indicate that photorespiration was relativelygreater at higher temperatures.     

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