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     

THE PHYSIOLOGICAL BASIS OF MORPHOLOGICAL POLARITY IN REGENERATION. I

1. In Bryophyllum calycinum two apical leaves suppress the shoot formation in all the dormant buds situated basally from the leaf; one apical leaf suppresses the shoot formation in the basal buds situated in the same half of the stem where the leaf is, and, if one-half of the petiole of such a leaf is removed, the growth of basal buds in one quadrant of the stem is suppressed. 2. This inhibitory influence of a leaf upon shoot formation in the basal part of a stem is diminished or disappears when the mass of the leaf is reduced below a certain limit. 3. The inhibitory influence of an apical leaf upon the growth of shoots in horizontally suspended stems is greater when the leaf is on the upper than when it is on the lower side of the stem. 4. All these facts suggest the possibility that the inhibitory influence of the leaf upon shoot formation is due to inhibitory substances secreted by the leaf and carried by the sap from the leaf towards the base of the stem. 5. An apical leaf accelerates root formation in the basal part of a stem and this accelerating effect increases with the mass of the leaf. 6. This inhibitory influence of a leaf upon shoot formation and the favoring influence upon root formation in the more basally situated parts of the stem is one of the factors determining the polar character of regeneration.     

THE PHYSIOLOGICAL BASIS OF MORPHOLOGICAL POLARITY IN REGENERATION. II

1. The experiments show that the mass of air roots formed in a stem increases with the mass of the leaf attached to the stem, though it has not been possible to establish an exact mathematical relation between the two masses, owing to unavoidable sources of error. 2. Darkened leaves do not increase the mass of roots formed. 3. In stems suspended horizontally air roots appear on the lower side of the stem, with the exception of the cut end where they usually appear around the whole circumference of the stem. When the lower half of a stem suspended horizontally is cut off, roots are formed on the upper side. It is shown by experiments on leaves suspended horizontally that the more rapidly growing roots and shoots on the lower side inhibit the root and shoot formation in the upper half of such a leaf; and likewise the more rapid formation of roots on the lower side of a horizontally suspended stem seems to account for the inhibition of root formation on the upper side of such a stem. Likewise the more rapid growth of shoots on the upper side of a stem suspended horizontally is likely to inhibit the growth of shoots on the lower side. 4. Each leaf contains in its axil a preformed bud capable of giving rise to a root, which never grows out in the normal stem on account of the inhibitory influence of the normal roots at the base of the plant. These dormant root buds are situated above (apically from) the dormant shoot bud. The apical root buds can be caused to develop into air roots when a piece of stem is cut out from a plant from which the leaves except those in the basal node of the piece are removed. The larger these basal leaves the better the experiments succeed. 5. These apical air roots grow out in a few days, while the roots at the basal end of the stem (which in our experiments dip into water) grow out about a week later. As soon as the basal roots grow out in water they cause the air roots in the more apical region of the stem to dry out and to disappear. 6. In addition to the basal roots, basal nodes have also an inhibitory effect on the growth of the dormant root buds in the apical region of a stem. This is indicated by the fact that a stem with one pair of leaves near the base will form apical air roots more readily when no node is situated on the stem basally from the leaf than if there is a node basally from the leaf.     

Nitrate Assimilation in Higher Plants with Special Reference to the Cocklebur (Xanthium pennsylvanicum Wallr.)

A soluble NADH-dependent nitrate reductase is described forthe shoot system of Xanthium. Young leaves and immature stemtissues contain high levels of the enzyme. They are relativelyrich in free amino acids and amides but store little free nitrate.The specific activity of the enzyme is lower in fully expandedleaves, although these leaves exhibit higher rates of fixationof carbon in photosynthesis than do younger leaves. Neithernitrate nor free amino acids accumulate in the mesophyll ofthe leaf. Older parts of the stem axis accumulate large amountsof soluble nitrogen, almost entirely as free nitrate. Reservesof nitrate in the shoot and root are rapidly depleted if nitrateis removed from the external medium. Nitrate reductase is apparently absent from roots of Xanthium.This finding is supported by analyses of bleeding sap from nitrate-fedplants which show that 95 per cent of the nitrogen exportedfrom roots is present as free nitrate. However, roots are capableof synthesizing and exporting large amounts of amino nitrogenif supplied with reduced nitrogen such as urea or ammonium. A scheme is presented summarizing the main features of the metabolismof nitrate in Xanthium and this is compared with the situationin nitrate-fed plants of the field pea (Pisum arvense L.), aspecies previously shown to be capable of reducing nitrate inits root system.     

Studies in the Nutrition of Apple Rootstocks: II. Effects of Concentration of Iron in the Nutrient Solution on Growth and Chlorophyll Content

Rooted one-year shoots were grown for one season by sprayingtheir roots with nutrient solution. Iron supplied as Fe-EDTAat four concentrations resulted in plants which were respectively(a) severely chlorotic, (b) mildly chlorotic, (c) dark greenand healthy (controls), and (d) dark green but with slight reductionin growth. Severely deficient plants showed 40–70 per cent reductionsin growth as measured by fresh weight, shoot length, diameterincrease, leaf area, net assimilation and relative growth-rates.Dry weights were reduced 70–80 per cent and of the totaldry-weight increment a greater proportion remained in the leaves,which had a lower dry weight and higher water content per unitarea. However, because the initial old stem formed a greaterproportion of the total dry weight, the leaf area ratio remainedabout 11 per cent lower than in the controls. Severely deficientplants had, per unit of chlorophyll, a higher dry-weight increaseand net assimilation rate than the controls. Mild deficiency caused 10–20 per cent reductions in growthand net assimilation rate; the leaf area ratio was normal. Possible mechanisms of the effects of low iron supply are discussed,while the small growth reduction at the highest Fe-EDTA concentrationis attributed to chelate toxicity     

THE ROLE OF RESERVES IN LEAVES,BRANCHES, STEMS,AND ROOTS ON SHOOT GROWTH OF RED PINE

By defoliation, girdling, and various combinations of these before the growing season began, the role of foods in old needles, branches, main stems, and roots on shoot growth of 8-yr-old red pine (Pinus resinosa) trees was studied in northern Wisconsin. Defoliation and girdling, alone or in combination, reduced shoot growth but defoliation reduced shoot dry weight more than it reduced shoot elongation. Reductions in shoot growth due to treatment were in the following decreasing order: branch girdling + needle removal > girdling at stem base + needle removal > needle removal > branch girdling > base girdling. The old needles were the major source of food for shoot growth and accounted for four-fifths or more of all shoot growth. The combined reserves in the branches, main stem, and roots accounted for less than 15% of shoot growth. The contribution of reserves from tissues other than old leaves was in the following order: branches > main stem > roots. Defoliation weakened apical dominance relations. In defoliated trees, many secondary axes elongated more than the terminal leader, and secondary axes in lower whorls often grew more than those in upper ones. The data suggest an important role of nutritional factors in correlative growth inhibition.     

Effect of Removal of a Part of the Root System on the Subsequent Growth of the Root and Shoot

Removal of up to 50 per cent. of the roots of barley and ryehas no effect on the growth-rate of the root which is the sameas in the intact plant. In contrast the growth-rate of the shootdecreases as more roots are removed. When more than 50 per cent.of the roots are removed, root growth declines but not so rapidlyas that of the shoot. Similar results are obtained by the removalof lateral roots of tomato but root growth begins to declinewhen 40 per cent. of the lateral roots are removed. The uptake of potassium by barley plants with proportions ofthe root system excised is closely proportional to the dry-matterincrease when the nutrient supply is not limiting. In conditionsof low nutrition the potassium uptake is less than the dry-matterincrease.     

Effects of Drought on Partitioning of Nitrogen in Two Wheat Varieties Differing in Drought-tolerance

A model was constructed to describe the translocation and partitioningof nitrogen on the seventh day after anthesis for well-wateredand droughted plants of two wheat varieties (Triticum aestivumL. cv. Warigal and Condor). The glasshouse-grown plants weredetillered so that a simplified model could be derived for themain stem. A 9-d drought treatment was imposed just after anthesisand this coincided with the period of endosperm cell divisionin the grains. Warigal, which had a higher grain yield thanCondor under drought, absorbed up to 15-times more nitrogenand translocated 1.5-fold more nitrogen to the shoot via thexylem. In both varieties, nitrogen redistributed from vegetativeorgans accounted for more than 60 per cent in control and 70per cent in droughted plants of the nitrogen needed for eargrowth. The net loss of nitrogen increased by 4-3 per cent inthe leaves, but decreased by 60 per cent in the stem under drought.Stem and roots appeared to play an important role in the nitrogeneconomy of droughted plants: less nitrogen was translocateddirectly to the grains from the senescing leaves and 40–60per cent more nitrogen was translocated to the roots. Nearlyall the nitrogen reaching the roots in the phloem was reloadedinto the xylem stream and translocated back to the shoot. Thetransfer of nitrogen through the stem was reduced under droughtand this resulted in a constant C:N ratio of the grains whichmay be important in the regulation of endosperm cell division. Triticum aestivum L., wheat, drought, nitrogen, senescence, translocation     

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.     

Jasmonic acid treatment to part of the root system is consistent with simulated leaf herbivory, diverting recently assimilated carbon towards untreated roots within an hour

It is known that shoot application of jasmonic acid (JA) leads to an increased carbon export from leaves to stem and roots, and that root treatment with JA inhibits root growth. Using the radioisotope ¹¹C, we measured JA effects on carbon partitioning in sterile, split-root, barley plants. JA applied to one root half reduced carbon partitioning to the JA-treated tissue within minutes, whereas the untreated side showed a corresponding – but slower – increase. This response was not observed when instead of applying JA, the sink strength of one root half was reduced by cooling it: there was no enhanced partitioning to the untreated roots. The slower response in the JA-untreated roots, and the difference between the effect of JA and temperature, suggest that root JA treatment caused transduction of a signal from the treated roots to the shoot, leading to an increase in carbon allocation from the leaves to the untreated root tissue, as was indeed observed 10 min after the shoot application of JA. This supports the hypothesis that the response of some plant species to both leaf and root herbivores may be the diversion of resources to safer locations.     

A Quantitative Analysis of the Uptake of Carbon and of the Supply of 14C-labelled Assimilates to Areas of Meristematic Growth in Lolium temulentum

A quantitative analysis of the ¹⁴C-labelled assimilate suppliedby leaves on the main shoot to terminal meristem, stem, tillers,and roots was conducted during parallel periods of reproductiveand vegetative development in Lolium temulentum. The initial rate of entry of carbon into the shoot varied withthe area and photosynthetic efficiency of the assimilating leaf.Subsequently, respiratory losses of carbon during translocationand incorporation of assimilate at the site of utilization alsovaried. The combined effect of these differences resulted inthe supply of recently assimilated carbon being twofold greaterin reproductive shoots than in vegetative shoots, while withinshoots the carbon supply of the youngest fully expanded leafranged from four-or five-fold greater than the oldest leaf inyoung shoots, to two-or three-fold greater in older shoots.In both reproductive and vegetative shoots, the two or threeyoungest leaves thus dominated the supply of carbon for meristematicgrowth. Meristematic tissue in expanding leaves and leaf primordia atthe terminal meristem of the vegetative shoot received 18–27per cent of the total shoot carbon. This meristem utilized aboutthe same proportion of shoot carbon when it developed into aninflorescence, indicating no major change in the level of meristematicactivity. The proportion of shoot carbon utilized in stem growthincreased as both reproductive and vegetative shoots aged; thisincreased meristematic activity in stem internodes was accompaniedby reduced export of carbon to roots, which received less than10 per cent of the shoot carbon when the experiments ended.The main shoot translocated 20–30 per cent of its recentlyassimilated carbon to developing and rooted tillers, which assinks for carbon were thus as important as the terminal meristemand stem. This outward flow of carbon continued relatively uncheckedwhen donor and receptor shoots developed inflorescences.     

Dipeptides and Amino Acids as Nitrogen Sources for the Callus of Atropa belladonna

Morphactin ester applied as a bark band to stems of Monterey pine seedlings (Pinus radiata D. Don) at first stimulated and then inhibited the rate of stem elongation over a 20-day period. The inhibition was caused by a direct action of morphactin on the shoot rather than by indirect action on the root system. ¹⁴C labelled morphactin applied to the bark accumulated in the shoot apices and the amount accumulated could be related to the inhibition of growth. More than 70% of labelled material isolated from roots and shoot apices was in the form of a metabolite similar to the carboxylic acid derivative of the morphactin ester. A preparation of ¹⁴C labelled acid derivative applied to the bark was translocated to the shoot apex and to the roots and inhibited stem elongation.     

Change in Electrophoretic Pattern of Ribonuclease during Dormancy and Growth of Bird-cherry Buds

A morphactin formulation containing methyl-2-chloro-9-hydroxyfluorene-9-carboxylate as the major component caused abnormal stem growth in Pinus radiata D. Don seedlings when applied as a bark band. The banded region swelled due to increased cell division in the periderm and cambium. The abnormal stem development in treated trees did not cause a localized girdling effect. The morphactin treatment inhibited shoot elongation and root growth. Treated trees exposed to ¹⁴CO₂ translocated significantly less assimilate to the roots than did control trees although fixation of ¹⁴CO₂ by the shoots was not reduced. Label from a bark band of ¹⁴C-morphactin was rapidly translocated to the roots indicating that direct inhibition of root growth by the inhibitor probably occurred.     

14C-Studies on Apple Trees III. The Influence of Season on Storage and Mobilization of Labelled Compounds

The present paper reports an attempt to elucidate the storage and mobilization processes in 1-year-old apple rootstocks by studying the ¹⁴C content of different parts of the tree following application of ¹⁴CO₂. The translocation of the ¹⁴C taken up from the leaves to other parts proceeds at the highest rate during the first few days after the application of ¹⁴C. The distribution in shoots, trunk and roots after application of ¹⁴C during May, July, August, and in part of September depends in particular upon the intensity of growth in the various parts. From the lime of the application of ¹⁴C until leaf-fall, 40–50 per cent of the ¹⁴C initially absorbed disappears from the tree. After exposure to ¹⁴C during October, and in part of September, a relatively large part of the ¹⁴C applied goes to the root. In this case there is a considerable reduction in the ¹⁴C-conlent from leaf-fall to the following spring after leafing, especially in the root, although relatively speaking reduction is also considerable in the bark of the parts above ground, and it is most pronounced in the methanol (80 %) soluble fraction. The reduction takes place primarily during spring, and comprises, after application during October, for the whole tree 20–25 per cent of the ¹⁴C initially absorbed. Only 13–17 per cent of this amount was recovered in the newly developed shoots and leaves in the following June.     

Comparison of radial increment and volume growth in stems and roots of Quercus petraea

Dendrochronological studies dealing with roots, stems and branches are very rare or often take the form of short notes. The difficulties of detecting rings and of quantifying the radial growth in roots have already been described for various species. In oak the anatomical root structure differs from stemwood. The roots are radial-porous or diffuse-porous, and there is often no clear distinction between individual rings. In our study visual and radiographic techniques were used to examine radial increment in roots of sessile oak (Quercus petraea L.) which was compared with radial growth in branches and along the stem. Coarse roots were cut from four 30- to 34-year-old trees that had been uprooted mechanically and disks were taken at different distances from the stem-root base. Ring widths were measured in the stem at height of 0.3 m, at breast height (1.3 m), beneath the crown, in branches of the crown, and in roots every 20 cm. The ring widths were cross-dated, and the heterogeneity of growth within a root and within the root system were analysed. Asymmetric growth frequently occurred in roots so that ovals, I-beam and T-beam shapes were developed. With the method used in our study the annual growth layers close to the central cylinder could be distinguished as well as beneath the bark. Pointer years were detected in all sections of the tree and permitted correction of ring widths in roots. Root system, stem and branch showed a basic similarity in their radial sequence of ring width. The annual biomass increment was weaker and more variable with several consecutive changes in the roots than in the stems. The root/shoot ratio reached a minimum rather early, beginning at the cambial age of 20 years. This revised version was published online in June 2006 with corrections to the Cover Date.     

The Influence of Moisture Stress on Liriodendron tulipifera L. Seedlings

A comparison of yellow poplar seedlings subjected to availablemoisture ranging from 100 per cent to 20 per cent of field capacityin two separate experiments (a soil study and a solution study)indicated that total dry matter production, stem height, totalleaf area and area of individual leaves, decreased with increasedmoisture stress. Differences in unit leaf rate (ULR) were primarilyresponsible for final plant size. Leaf area: weight ratio wasaffected by the method of induced stress. As stress increasedthe ratio increased in the soil study but decreased in the solutionstudy. The number of leaves produced was unaffected by stress,except at the most severe stresses. The number of senescentleaves increased as moisture stress increased. The allocationof new growth was unaffected by either moisture stress or timeand decreased in order of root, foliage, stem for the soil grownplants and foliage, root, stem for the solution grown plants.ULR and relative growth rate (RGR) were reduced by both moisturestress and time. A growth simulation of the soil-grown plantsbased on results from the solution moisture stress experimentpredicted the final plant dry weight within 12 per cent andleaf area within 7 per cent of the actual values.     

Distribution of assimilate during the flush cycle of growth in Theobroma cacao L.

The growth of the shoot and roots of seedling plants of cocoa (Theobroma cacao L.) under constant glasshouse conditions showed a rhythmic cycle, with the maximum growth stages of each alternating in a regular sequence. When the growth cycle of the shoot was upset by removing all new leaves immediately after unfolding, the roots showed a high constant growth rate during this period, suggesting that normally the rapidly expanding leaves exert an inhibitory influence on the roots. Conversely removal of portions of the root delayed the production of new leaves in the shoot. The level of soluble and starch carbohydrate in the mature leaves, roots and stem declined during the period of expansion of the flush leaves, but accumulated again at the end of the leaf expansion stage. It is likely that this reserve carbohydrate was remobilised and translocated to the flush leaves during their period of expansion. A large proportion of newly formed photoassimilate, as shown by the distribution of ¹⁴C radioactivity from different source leaves, was also translocated to the young leaves during expansion. The large sink created by these leaves may cause photoassimilate and reserve carbohydrate to be diverted from the roots, thereby inhibiting root growth during the stage of leaf expansion. It is suggested that the rhythmic leaf production at the apex may control the growth cycle of the roots.     

温州蜜柑萎缩病毒和柑桔衰退病毒在苗木各部位分布的全年分析

１９９１年９月至１９９２年８月逐月采样,采用ＤＡＳ－ＥＬＩＳＡ法对温州蜜柑萎缩病毒（ＳＤＶ）在温州蜜柑苗木各部位及对柑桔衰退病毒（ＣＴＶ）在甜橙苗木各部位的分布进行了全年分析。结果表明ＳＤＶ在老叶、老枝皮中全年均检测不到;在根中仅在７、８月份可检出有少显病毒存在;在嫩叶中３、４、５、６、９、１０月均可检出且浓度较高,而７、８月在嫩叶中检测不出,在嫩枝皮中４、５、６、７、１０月均可检出而７月份浓度有较大幅度降低。这说明冷凉温度下的嫩组织适立于ＳＤＶ的繁殖,最佳检测时期为春梢和秋梢的嫩梢期,最佳检测部位为幼嫩的叶和嫩枝皮。对ＣＴＶ在甜橙苗木各部位分布检测结果表明各部位全年均可检测出带毒且含量均较高,总的变幅不大,冬季和夏季含量有所下降,在老叶中变幅稍大些,这说明甜橙苗木各个部位均适宜于采样且全年均可进行分析。     

Phosphorus nutrition of jarrah (Eucalyptus marginata) seedlings

Summary Effects of calcium phosphate supply on plant dry matter and phosphorus concentrations of parts of jarrah (Eucalyptus marginata) seedlings grown in a lateritic topsoil from the jarrah forest were examined in two glasshouse trials. Phosphorus deficiency depressed root and shoot dry weights and severely deficient leaves were smal and purple with prominent red major veins. Phosphorus deficiency severely reduced stem phosphorus levels (0.5% to 0.02%, experiment 1). Phosphorus concentrations were higher in bark than wood and the amount of phosphorus in the bark was sensitive to stem age and phosphate supply. Phosphorus adequate plants had bark phosphorus concentrations in the range 0.2–0.9% compared to <0.1% in deficient plants (experiment 2). Jarrah leaves accumulated dry matter up to 80 days after expansion and some leaves exported phosphorus during this period. Bark analysis may therefore be preferable to leaf analysis for detecting phosphorus deficiency in this species.     

Role of roots in regulating the growth rate and cytokinin content in leaves

The role of roots in the enhancement of cytokinin content and leaf growth of Phaseolus vulgaris plants after decapitation and partial defoliation was investigated. Partial excision of the roots of plants which were decapitated above the primary leaf node resulted in a reduction of leaf growth and soluble proteins accumulation in the primary leaves. Roots excision was done at time of decapitation and repeated 8 days later. Endogenous cytokinins, known to be involved in enhancing shoot growth, accumulated in the leaves and stems of decapitated and partially defoliated plants. Lower levels of cytokinins were detected in the leaves of decapitated plants with only a partial root system. The level of cytokinins in the roots of decapitated plants was reduced by partial root excision. The growth and accumulation of cytokinins in leaves were, however, not totally suppressed by removing a large proportion of the roots. At the commencement of the experiment the stem had a higher cytokinin content than both the leaves and roots. This suggests that the stem could be an alternative source of cytokinins to the leaves. The cytokinin complement in the leaves of decapitated plants is not identical to that in the roots. It appears that cytokinins supplied by the roots are metabolized in the leaves, or that alternatively certain cytokinins are synthesized in the leaves themselves.