Distribution of labelled assimilates within a young apple tree after supplying14CO2to a leaf or shoot

Labelled carbon dioxide was supplied for 22 hrs to a leaf of the leader or to the lateral shoot in two-year-old apple seedlings. The distribution of radioactive assimilates within the plant following this treatment was investigated by using radioautography. The transport of labelled assimilates from the young leaf of the leader was very meagre and affected only parts of the stem and the leaves situated in the close vicinity of the treated leaf. The¹⁴C-labelled assimilates from the mature leaf of the leader were transported in a considerable amount to the apex and to the other leaves of the leader. They were also found in an appreciable amount in the stem and the roots, as well as in some lateral shoots. After supplying¹⁴CO₂ to the lateral shoot remarkable transport of labelled assimilates was observed. Radioactivity was detected in the tip and in the youngest leaves of the leader, as well as in the roots. Their path in the stem was studied by dissecting the plant and examining the cross section from each internode. This method revealed that the assimilates from the treated leaf or shoot were transported downward only on one side of the stem in a helical pattern. The lateral shoots situated on the radioactive side of the stem were also labelled, whereas those situated on the opposite (non-radioactive) side were not labelled.     

Photosynthesis and distribution of photosynthates in apple shoots treated by pinching and bark ringing

Individual leaves on apple shoots treated by pinching and/or ringing were supplied with¹⁴CC₂ and the rate of photosynthesis and the pattern of photosynthate distribution was studied. The ringing of the shoots as well as the ringing and pinching done together reduces the rate of photosynthesis almost by half. Pinching itself had no effect on the rate of photosynthesis. Removing the main sink — the growing shoot apex — by pinching leads to the accumulation of photosynthates in the shoot, mainly in the first internode below the pinching point. Accumulated assimilates were soon used up by the axillary meristems which were stimulated to grow by pinching. Very small amounts of assimilates (about 1 %) were translocated to adjacent shoots which were supplied by their own leaves. When adjacent shoots were defoliated, more assimilates were translocated there (nearly 8%), however, a strong tendency to accumulate the assimilates in the internode below the pinching point was observed. Assimilates are not located in matured leaves. The presence of an active sink located near the donor stimulates the mobilization of assimilates present in the donor leaf.     

Translocation of some assimilates from the sink to the donor in apple tree

The shoot apex or fruitlets of Jonathan apple trees grafted on M IX rootstock and grown in pots in a greenhouse were exposed to¹⁴CO₂ in an assimilation chamber. The translocation of¹⁴C-labelled assimilates from treated organs to other parts of the plant was studied. It was found that a very small amount of¹⁴C-labelled compounds was translocated from the shoot apex and very young fruitlets to the shoot stem. Preliminary chromatographic studies show that the chemical composition of the labelled substances detected below assimilation chamber differs profoundly from that of those remaining in the supplied leaves. The results support the view that there exists a translocation of some substances, possibly regulators from the sink to the donor.     

Seasonal differences in the pattern of assimilate movement in branches of Coffea arabica L

At Ruiru, Kenya, ¹⁴CO₂ was fed to single leaves at different distances from the apex on branches of mature fruiting and non-fruiting trees of Coffea arabica on six occasions from the Short Rains (November) 1966 to the Long Rains (April-May) 1967. The location of ¹⁴C-labelled assimilates in the treated branches was determined 26 h later by autoradiography. The direction of movement of labelled assimilates indicated large seasonal differences in the relative sink strengths of the shoot tip, fruit and trunk-root systems. On vegetative trees the sink strength of trunk-roots was much smaller, as compared with the shoots, at the beginning of the Long Rains than at the end of the previous Short Rains or in the intervening dry season. Assimilate use by growing fruits did not alter the pattern of distribution of assimilates to the other sinks at the end of the Short Rains, but it did restrict assimilate movement to both shoot tips and trunk-roots at the_ beginning of the Long Rains. In the dry season, virtually all assimilates were utilized by growing fruits when these were present. Vegetative secondary shoots provided assimilates to growing fruits and trunk-roots at the end of the Short Rains and in the dry season. Some practical implications are noted.     

The Translocation of 14C-Labelled Assimilates in Tomato Plants Infected with Alternaria Solani (Ell. and Mart.) Jones and Grout

A quantitative study was made of the effect of infection bya facultative parasite, Alternaria solani, on the translocationof ¹⁴C-labelled assimilates in the tomato plant. In a plantwith a single diseased leaf, this leaf retained a greater proportionof its assimilated radiocarbon during the early stages of infection.The export of assimilates from a diseased leaf when diseasedevelopment was purposely retarded and was found to be similarto that from a control leaf. At a late stage in the disease,where there was extensive chlorosis and necrosis, export fromdiseased leaves was generally increased and the distributionof the translocated assimilates was altered. Non-infected leaveswere at first unaffected by the disease, but later there wasincreased contribution to the actively growing regions of theplant. In some cases there was a slight increase in total exportfrom healthy leaves but, more generally, a change in the distributionof assimilates occurred. In particular, there was a consistentincrease in activity in the roots. This response to infectionwas not related to the amount of disease present. No evidencewas obtained for an increased import into infected leaves fromnon-infected leaves at any stage of the disease.     

Uptake and translocation of paclobutrazol by shoots of M.26 apple rootstock

When ¹⁴C-paclobutrazol, a gibberellin synthesis inhibitor, was applied to different parts of actively-growing M.26 apple rootstock shoots it was translocated acropetally when applied to the young stem and, to a lesser extent, from the youngest unrolled leaf. Paclobutrazol was not translocated out of leaf laminae, shoot tips or from one-year-old wood but translocation occurred out of a treated petiole into the attached leaf. No basipetal translocation was detected. This translocation pattern suggested movement through the xylem.Localised application of paclobutrazol caused a reduction in shoot extension and leaf production when the young stem or shoot tip were treated; the effect decreased as older parts of the stem were treated. Treatment of laminae or petioles had only a slight effect on shoot extension and treatment of one-year-old wood was ineffective. Combined treatment of the shoot tip plus young stem was similar in effect to treatment of the complete shoot.It is suggested that paclobutrazol exerts its effects on shoot growth by inhibiting gibberellin biosynthesis in the shoot tip and the expanding leaves.The findings contribute to an understanding of the requirements for efficient orchard application of foliar sprays of paclobutrazol.     

Independent translocation of 14C-labelled assimilates and of the floral stimulus in Lolium temulentum

Summary It is widely accepted that the floral stimulus produced in leaves is carried to the shoot apex passively in the phloem with the assimilate stream. Three kinds of evidence presented here suggest that the floral stimulus moves independently of the assimilates.Simultaneous determination of the velocities of translocation out of the seventh leaf blade, in comparable plants under the same conditions, yielded estimates of 1–2.4 cm/hr for the floral stimulus, and 77–105 cm/hr for ¹⁴C-labelled assimilates.The effect of the size of the seventh leaf on its ability to export assimilates or to initiate flowering was quite different. Leaves with only 14–26% of their final blade area emerged exported little assimilate, yet were highly active in inducing flowering.The effect of DCMU applications at a range of concentrations on the translocation of assimilates was quite different from their effect on the flowering response.     

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.     

Translocation and degradation of injected uniconazole in apple during a 4-month growing period

One-year-old ‘Golden Delicious’ apple trees grafted onto MM 106 rootstocks were injected in the rootstock stem (shank) with¹⁴C-uniconazole to determine the extent to which uniconazole is translocated and degraded over the length of an average growing season. In 4 months, 16% of recovered¹⁴C-activity was translocated to the new shoots. Most of the¹⁴C-activity remained in the rootstock. Chromatographic evaluation of shoot extracts demonstrated that the¹⁴C-activity associated with uniconazole decreased 49% in 4 months. However, shoot growth was still inhibited which suggests that the amount of uniconazole that was degraded did not interfere with the inhibition of gibberellin biosynthesis, probably due to the continuous translocation of uniconazole that occurred.     

Floral stimulus movement in perilla and flower inhibition caused by noninduced leaves

Photoperiodic control of flowering in the short day plant Perilla involves the transmission of a floral stimulus from induced leaves to the shoot apex. We have studied the basipetal movement of this stimulus and of ¹⁴C-labeled assimilates in plants with an induced leaf (donor) grafted into the uppermost internode of a vegetative plant in which the axillary shoots at various nodes along the stem function as receptors.     

Branch bending affected floral bud development and nutrient accumulation in shoot terminals of ‘Fuji’ and ‘Gala’ apples

Branch bending has been practiced for decades in China to induce flower buds in ‘Fuji’ apple. However, the optimum bending angle is yet to be elucidated. The main objectives of this study were to compare the effect of branch bending angles (70°, 90° and 110°) on the flowering and nutrient accumulation of 1-year-old shoots of ‘Fuji’ and ‘Gala’ apples and to determine the optimum branch bending angle for each cultivar. In both cultivars, the production of spurs and terminal flower buds, and the total sugar concentration and the carbon-to-nitrogen (C/N) ratio in the shoot terminals increased, whereas the N concentration decreased with increasing bending angles. The nutrient concentration was significantly higher in spurs than in medium and long shoots. The distinction between the changing patterns of C and N concentrations in the bent shoots during the growing season in our study suggested the competition of these two nutrients caused by vegetative and reproductive growth at different growing times. In ‘Fuji’ apple, the proportion of flowering buds appeared to increase more rapidly with the increase of bending angle from 70° to 110° than that in ‘Gala’ apple, and particularly a higher proportion of spurs was observed on ‘Fuji’ branches bent at a larger angle. The increase in the total sugar concentration and the C/N ratio in the shoot terminals of the bent branches might be involved in inducing floral buds after bending. The optimum bending angle was about 90° for ‘Gala’ apple and 110° for ‘Fuji’ apple, respectively. Bending could help farmers to reduce the severity of biennial fruiting in ‘Fuji’ apple.     

Effect of Ophiobolus graminis infection on the assimilation and distribution of 14C in wheat

The uptake of ¹⁴C and movement of ¹⁴C-labelled assimilates in wheat plants inoculated with Ophiobolus graminis was examined following exposure of the second youngest leaf to ¹⁴CO₂. Autoradiographs of plants with infected seminal roots showed that assimilates were not translocated past the sites of root infection but accumulated in the undamaged portions of infected root systems, in particular the developing crown roots. There was no evidence that assimilates accumulated in the vicinity of O. graminis lesions. The net assimilation of ¹⁴CO₂ by wheat plants over a 5 h feeding period was not significantly affected by O. graminis infection. However, infection reduced the amount of ¹⁴C lost through respiration. Infection delayed the transfer of labelled assimilates from the fed leaf to the remainder of the plant but increased the proportion translocated to the roots. The latter effect was not apparent when infected plants were continuously irrigated during, and for 20 h following, the feeding period.     

Transport of exogenous auxin in two-branched dwarf pea seedlings (Pisum sativum L.)

Dwarf pea plants bearing two cotyledonary shoots were obtained by removing the epicotyl shortly after germination, and the patterns of distribution of ¹⁴C in these plants was investigated following the application of [¹⁴C]IAA to the apex of one shoot. Basipetal transport to the root system occurred, but in none of the experiments was ¹⁴C ever detected in the unlabelled shoot even after transport periods of up to 48 h. This was true both of plants with two equal growing shoots and of plants in which one shoot had become correlatively inhibited by the other, and in the latter case applied whether the dominant or subordinate shoot was labelled. In contrast, when [¹⁴C]IAA was applied to a mature foliage leaf of one shoot transfer of ¹⁴C to the other shoot took place, although the amount transported was always low. Transport of ¹⁴C from the apex of a subordinate shoot on plants bearing one growing and one inhibited shoot was severely restricted compared with the transport from the dominant shoot apex, and in some individual plants no transport at all was detected. Removal of the dominant shoot apex rapidly restored the capacity of the subordinate shoot to transport apically-applied [¹⁴C]IAA, and at the same time led to rapid cambial development and secondary vascular differentiation in the previously inhibited shoot. Applications of 1% unlabelled IAA in lanolin to the decapitated dominant shoot maintained the inhibition of cambial development in the subordinate shoot and its reduced capacity for auxin transport. These results are discussed in relation to the polarity of auxin transport in intact plants and the mechanism of correlative inhibition.Abbreviations IAA Indol-3-yl-acetic acid - TIBA 2,3,5-triiodobenzoic acid - 2,4D 2,4-dichlorophenoxyacetic acid - IAAsp Indol-3-yl-acetyl aspartic acid     

Translocation of 14C-Assimilates in Roses

Movement of ¹⁴C-assimilates from young and mature leaves to young rose shoots (Rosa hybrida cv. Marimba) was examined in two developmental stages. In the first stage after bud breaking the young shoot, especially its tip, depends for its supply of assimilates mainly on the mature foliage. At this stage young leaves are powerful sinks and retain 97% of their own photosynthates. The translocated 3% move mainly to the roots. At a later stage, just after the appearance of the flower bud, most of the leaves on the shoot become a source. The upper leaves supply assimilates to the flower bud and to the upper part of the stem. The ¹⁴C-assimilates from the lower leaves move in two directions, the larger part being directed downward.     

The utilization of recently assimilated carbon in graminaceous plants

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

Effect of assimilate utilization on photosynthetic rate in wheat

Summary Two weeks after anthesis, when the grain is filling rapidly, the rate of photosynthesis by flag leaves of wheat cv. Gabo was between 20 and 30 mg CO₂ dm^-2 leaf surface hour^-1 under the conditions used. About 45% of flag-leaf assimilates were translocated to the ear, and only about 12% to the roots and young shoots.On removing the ear, net photosynthesis by the flag leaves was reduced by about 50% within 3–15 hours, and there was a marked reduction in the outflow of ¹⁴C-labelled assimilates from the flag leaves.Subsequent darkening of all other leaves on plants without ears led to recovery of flag-leaf photosynthesis, as measured by gas analysis and ¹⁴CO₂ fixation, and to increased translocation of assimilates to the roots and young shoots. Minor changes in the rates of dark respiration accompanied these major, reversible changes in photosynthetic rate.After more than a week in continuous, high-intensity light, the rate of photosynthesis by flag leaves of intact plants had fallen considerably, but could be restored again by a period in darkness, or by inhibiting photosynthesis in the ears by spraying them with DCMU. The inhibition of ear photosynthesis increased translocation of labelled assimilates from the flag leaf to the ears, without affecting leaf sugar levels.The application of TIBA to the culm below the ear inhibited auxin movement throught the culm, but had no influence on flag-leaf photosynthesis.These results suggest that, at least in this system, photosynthesis by the flag leaf is regulated directly by the demand for assimilates from the flag leaf and not indirectly through action in the leaf of auxins produced by the sink organs.     

14C-Studies on Apple Trees. II. Distribution of Photosynthates from Top and Base Leaves from Extension Shoots

The photosynthates from leaves from extension shoots may be used either for new growth of the shoot itself, or for growth in other parts of the tree. An attempt has been made to elucidate this problem by determining the content of ¹⁴C in long shoots to which ¹⁴CO₂ was applied either through fully developed leaves at the base, or through very young leaves at the apex. In the case of ¹⁴C application at the base, 80 per cent or more of the ¹⁴C initially taken up disappears from the shoot, and only a very minor part is translocated upwards in the shoot. When the leaves at the apex are exposed, on the other hand, 80 per cent of the ¹⁴C absorbed is retained in the leaves and shoot components treated as long as there is still considerable terminal growth taking place, although a small percentage is deposited in the lower parts of the shoot. At the same time, a much higher proportion is incorporated into methanol-insoluble components. As terminal growth decreases, a larger proportion of the ¹⁴C activity of the apex leaves also disappears from the shoot. The distribution of activity between the sorbitol, sucrose, glucose and fructose fractions was not significantly different in young and in fully developed leaves. The ¹⁴C labelling in the sugar fraction was highest for sorbitol, then sucrose, but decreases with time compared to glucose and fructose.     

Phloem transport in Picea abies (L.) Karst. in mid-winter

Summary Translocation of ¹⁴C assimilates was studied on four different transport systems of Picea abies branches after induced activation in January. ¹⁴CO₂ assimilation of terminal shoots for 48 h at 25° C resulted in phloem loading and basipetal transport of ¹⁴C photosynthate into the following, older shoot generations. ¹⁴C import was enhanced, when these older shoot generations were kept in the dark. Microautoradiographs of the labelled terminal shoots showed that ¹⁴C assimilates were exported from needles via sieve elements of the leaf traces and loaded into the latest increment of the axial secondary phloem. No ¹⁴C label appeared in the obliterated sieve cells or in the tracheids. In addition, ¹⁴C photosynthate accumulated densely in the chlorophyllous cells of the cortex and in cells of the resin ducts, indicating certain sink activity. In the darkened 2-year-old shoot, imported ¹⁴C photosynthate was concentrated in the functional secondary phloem, while some ¹⁴C label was unloaded into the latest xylem increment. When 6-year-old shoots were exposed to ¹⁴CO₂ for 48 h in the light, ¹⁴C assimilates accumulated in the phloem of the leaf trace and in the latest increment of the axial secondary phloem. However, a substantial amount of radioactivity was unloaded into ray cells and phloem parenchyma cells. Thus, the presence of functioning phloem in needles and twigs of P. abies during winter allows long-distance translocation and radial distribution of assimilates according to existing source-sink relations.     

Sectorial Patterns in Leaves on Fruit Tree Shoots Produced by Radioactive Assimilates and Solutions

In vigorously growing shoots of apple and plum ¹⁴C-assimilateswere translocated from a ‘fed’ leaf to particularsectors of other leaves in a distribution pattern associatedwith the phyllotaxis; the same sectorial and distribution patternswere produced by ³²P phosphate solution taken into the shootthrough a cut petiole. The frequency with which a given sectorialpattern occurred at a particular position on the phyllotacticspiral was ascertained. Such patterns were not observed abovethe third rolled leaf in the apple shoot apex. Killing the phloem in the petiole prevented egress of labelledassimilate but not of ³²P solution. Barkringing above the sourceleaf reduced, but did not completely prevent, assimilate movementup the stem, suggesting some translocation in the xylem. Distribution of label from ⁴⁵CaCl₂, ⁸⁶RbCl and [³H]asparagine,incorporated through cut petioles, did not follow the same patternas label from ³²P solutions. Malus pumila Mill., apple, Prunus domestica L., Prunus insititia. L., leaf plum, patterns, transport of radioisotopes, vascular phyllotaxis     

Retention and allocation of <Superscript>14</Superscript>C assimilates by maintenance leaves and harvest index of tea (<Emphasis Type="Italic">Camellia sinensis</Emphasis> L.)

Partitioning of ¹⁴C-labelled photosynthates to various parts of un-pruned tea clones TV1 and TV25 was assessed in vivo by exposing maintenance leaves to ¹⁴CO₂ at monthly intervals throughout the year. The plants from shoot apex to root tip were divided into twelve components to assess the allocation and retention of ¹⁴C-photosynthates by the maintenance foliage. Out of the total photosynthates produced by the maintenance leaves, only 11.08 % was allocated to the commercially useful harvestable two and a bud shoots which is accepted as the harvest index of tea. The photosynthetically active maintenance leaves retained 19.05 % while 24.56 % was distributed to the branches. The bottom and the top parts of the trunk utilized 7.44 and 7.21 %, respectively. The thick roots at the base of the trunk, medium sized roots, pencil size roots, and feeder roots imported 7.28, 7.72, 7.65, and 8.01 % of ¹⁴C assimilates, respectively. Except retention by leaves, all the plant parts of vigorous clone TV25 required higher percentage of assimilates than TV1. The mean quantities of net photosynthates utilized by the stem and the roots were 69.37 and 30.63 %, respectively.