Water Transfer in an Alfalfa/Maize Association : Survival of Maize during Drought

We investigated the possibility of interspecific water transfer in an alfalfa (Medicago sativa L.) and maize (Zea mays L.) association. An alfalfa plant was grown through two vertically stacked plastic tubes. A 5 centimeter air gap between tubes was bridged by alfalfa roots. Five-week old maize plants with roots confined to the top tube were not watered, while associated alfalfa roots had free access to water in the bottom tube (the −/+ treatment). Additional treatments included: top and bottom tubes watered (+/+), top and bottom tubes droughted (−/−), and top tube droughted after removal of alfalfa root bridges and routine removal of alfalfa tillers (−^*). Predawn leaf water potential of maize in the −/+ treatment fell to −1.5 megapascals 13 days after the start of drought; thereafter, predawn and midday potentials were maintained near −1.9 megapascals. Leaf water potentials of maize in the −/− and −^* treatments declined steadily; all plants in these treatments were completely desiccated before day 50. High levels of tritium activity were detected in water extracted from both alfalfa and maize leaves after ³H₂O was injected into the bottom −/+ tube at day 70 or later. Maize in the −/+ treatment was able to survive an otherwise lethal period of drought by utilizing water lost by alfalfa roots.     

Phytophthora cryptogea root rot of tomato in rockwool nutrient culture:

The effect of root temperature on growth and yield of rockwool-grown tomato plants infected with Phytophthora cryptogea was investigated. Measurements of shoot and root growth were taken at high (25^oC) and low (15^oC) root temperatures during the generative phase of growth. The growth of roots of healthy and P. cryptogea-infected tomato plants in rockwool blocks was higher in plants grown with roots at 25^oC than at 15^oC after 60 days and a similar effect was found in slabs after 98 days. Under sub-optimal conditions for growth the disease became severe when root temperatures were low. Growth of roots was greatest when roots were maintained at a high temperature in combination with an ambient air temperature of c. 15^oC and the response was greater in cv. Counter than cvs Calypso and Marathon. Water-soluble carbohydrates of roots were higher in those produced in blocks than slabs and were reduced by infection compared to healthy plants with roots at 15^oC and 25^oC. Reduced transpiration rates were found 17 days after inoculation in symptomless plants grown at a root temperature of 25^oC. Infection, regardless of the temperature of the roots or cultivar, led to reduced stem growth. The plants grown at 25^oC were taller than those with a root temperature of 15^oC. After 9 wk of harvest, the cumulative fruit yields in infected cvs Counter and Calypso grown at 25^oC were comparable to that in healthy plants grown at either temperature and cumulative fruit numbers followed a similar pattern. High root temperatures led to delayed fruit ripening between weeks 3–10 and a larger number of unripe fruit. The weight of unripe fruit from infected plants grown at 25^oC at the terminal harvest was higher than from healthy plants with roots maintained at 15^oC.     

Enhanced N-Transfer from a Soybean to Maize by Vesicular Arbuscular Mycorrhizal (VAM) Fungi

Using a split-root technique, roots of soybean plants were divided between two pots. In one of the two pots, two maize plants were grown and half of those pots were inoculated with the vesicular arbuscular mycorrhizal (VAM) fungus, Glomus fasciculatus. Fifty-two days after planting, ¹⁵N-labeled ammonium sulfate was applied to the pots which contained only soybean roots. Forty-eight hours after application, significantly higher values for atom per cent ¹⁵N excess were found in roots and leaves of VAM-infected maize plants as compared with the non-VAM-infected maize plants. Results indicated that VAM fungi did enhance N transfer from one plant to another.     

Iron-Stress Induced Redox Activity in Tomato (Lycopersicum esculentum Mill.) Is Localized on the Plasma Membrane

Tomato plants (Lycopersicum esculentum Mill.) were grown for 21-days in a complete hydroponic nutrient solution including Fe³⁺-ethylenediamine-di(o-hydroxyphenylacetate) and subsequently switched to nutrient solution withholding Fe for 8 days to induce Fe stress. The roots of Fe-stressed plants reduced chelated Fe at rates sevenfold higher than roots of plants grown under Fe-sufficient conditions. The response in intact Fe-deficient roots was localized to root hairs, which developed on secondary roots during the period of Fe stress. Plasma membranes (PM) isolated by aqueous two-phase partitioning from tomato roots grown under Fe stress exhibited a 94% increase in rates of NADH-dependent Fe³⁺-citrate reduction compared to PM isolated from roots of Fe-sufficient plants. Optimal detection of the reductase activity required the presence of detergent indicating structural latency. In contrast, NADPH-dependent Fe³⁺-citrate reduction was not significantly different in root PM isolated from Fe-deficient versus Fe-sufficient plants and proceeded at substantially lower rates than NADH-dependent reduction. Mg²⁺-ATPase activity was increased 22% in PM from roots of Fe-deficient plants compared to PM isolated from roots of Fe-sufficient plants. The results localized the increase in Fe reductase activity in roots grown under Fe stress to the PM.     

Arbuscular Mycorrhizal Colonization Alters Subcellular Distribution and Chemical Forms of Cadmium in Medicago sativa L. and Resists Cadmium Toxicity

Some plants can tolerate and even detoxify soils contaminated with heavy metals. This detoxification ability may depend on what chemical forms of metals are taken up by plants and how the plants distribute the toxins in their tissues. This, in turn, may have an important impact on phytoremediation. We investigated the impact of arbuscular mycorrhizal (AM) fungus, Glomus intraradices, on the subcellular distribution and chemical forms of cadmium (Cd) in alfalfa (Medicago sativa L.) that were grown in Cd-added soils. The fungus significantly colonized alfalfa roots by day 25 after planting. Colonization of alfalfa by G. intraradices in soils contaminated with Cd ranged from 17% to 69% after 25–60 days and then decreased to 43%. The biomass of plant shoots with AM fungi showed significant 1.7-fold increases compared to no AM fungi addition under the treatment of 20 mg·kg⁻¹ Cd. Concentrations of Cd in the shoots of alfalfa under 0.5, 5, and 20 mg·kg⁻¹ Cd without AM fungal inoculation are 1.87, 2.92, and 2.38 times higher, respectively, than those of fungi-inoculated plants. Fungal inoculation increased Cd (37.2–80.5%) in the cell walls of roots and shoots and decreased in membranes after 80 days of incubation compared to untreated plants. The proportion of the inactive forms of Cd in roots was higher in fungi-treated plants than in controls. Furthermore, although fungi-treated plants had less overall Cd in subcellular fragments in shoots, they had more inactive Cd in shoots than did control plants. These results provide a basis for further research on plant-microbe symbioses in soils contaminated with heavy metals, which may potentially help us develop management regimes for phytoremediation.     

The symbiotic recapture of nitrogen from dead mycorrhizal and non-mycorrhizal roots of tomato plants

Aims

The aim was to quantify the nitrogen (N) transferred via the extra-radical mycelium of the arbuscular mycorrhizal fungus Glomus intraradices from both a dead host and a dead non-host donor root to a receiver tomato plant. The effect of a physical disruption of the soil containing donor plant roots and fungal mycelium on the effectiveness of N transfer was also examined.

Methods

The root systems of the donor (wild type tomato plants or the mycorrhiza-defective rmc mutant tomato) and the receiver plants were separated by a 30 μm mesh, penetrable by hyphae but not by the roots. Both donor genotypes produced a similar quantity of biomass and had a similar nutrient status. Two weeks after the supply of ¹⁵?N to a split-root part of donor plants, the shoots were removed to kill the plants. The quantity of N transferred from the dead roots into the receiver plants was measured after a further 2 weeks.

Results

Up to 10.6 % of donor-root ¹⁵N was recovered in the receiver plants when inoculated with the arbuscular mycorrhizal fungus (AMF). The quantity of ¹⁵N derived from the mycorrhizal wild type roots clearly exceeded that from the only weakly surface-colonised rmc roots. Hyphal length in the donor rmc root compartments was only about half that in the wild type compartments. The disruption of the soil led to a significantly increased AMF-mediated transfer of N to the receiver plants.

Conclusions

The transfer of N from dead roots can be enhanced by AMF, especially when the donor roots have been formerly colonised by AMF. The transfer can be further increased with higher hyphae length densities, and the present data also suggest that a direct link between receiver mycelium and internal fungal structures in dead roots may in addition facilitate N transfer. The mechanical disruption of soil containing dead roots may increase the subsequent availability of nutrients, thus promoting mycorrhizal N uptake. When associated with a living plant, the external mycelium of G. intraradices is readily able to re-establish itself in the soil following disruption and functions as a transfer vessel.     

Biosynthesis of the tigloyl esters of Datura: Cis-trans isomerism

Datura innoxia plants were wick fed with angelic acid-[1-¹⁴C] and l-isoleucine-[U-¹⁴C] to act as a positive control. After 7 days the root alkaloids 3α-tigloyloxytropane, 3α,6β-ditigloyloxytropane, and 3α,6β-ditigloyloxytropan-7β-ol were isolated and it was determined that angelic acid is not a precursor for the tigloyl moiety of these alkaloids. Tiglic acid-[1-¹⁴C] which was fed via the roots to hydroponic cultures of Datura innoxia, was incorporated to a considerable degree after 8 days.     

Effects of Oxime Carbamate Nematicides on Development of Heterodera schachtii on Sugarbeet

Treatment of sugarbeet, Beta vulgaris L., with aldicarb, aldicarb sulfoxide, or aldicarb sulfone 10 days after plants were inoculated with Heterodera schachtii prevented development of the nematode, but second-stage larvae penetrated the roots. These chemicals had no measurable effects on nematodes in plants treated 15 days after inoculation. The tests established that soil treatments of aldicarb are directly or indirectly lethal to larvae developing within roots of sugarbeet. Heterodera schachtii failed to develop on root slices of red table beet grown in soil treated with aldicarb or aldicarb sulfoxide. Similar treatment of plants with aldicarb sulfone or oxamyl did not affect subsequent development of H. schachtii on root slices of treated plants.     

Influence of inorganic nitrogen sources on K+/Na+ homeostasis and salt tolerance in sorghum plants

This work aimed to study the regulation of K⁺/Na⁺ homeostasis and the physiological responses of salt-treated sorghum plants [Sorghum bicolor (L.) Moench] grown with different inorganic nitrogen (N) sources. Four days after sowing (DAS), the plants were transferred to complete nutrient solutions containing 0.75 mM K⁺ and 5 mM N, supplied as either NO₃ ^? or NH₄ ⁺. Twelve DAS, the plants were subjected to salt stress with 75 mM NaCl, which was applied in two doses of 37.5 mM. The plants were harvested on the third and seventh days after the exposure to NaCl. Under the salt stress conditions, the reduction of K⁺ concentrations in the shoot and roots was higher in the culture with NO₃ ^? than with NH₄ ⁺. However, the more conspicuous effect of N was on the Na⁺ accumulation, which was severely limited in the presence of NH₄ ⁺. This ionic regulation had a positive influence on the K⁺/Na⁺ ratio and the selective absorption and transport of K⁺ in the plants grown with NH₄ ⁺. Under control and salt stress conditions, higher accumulation of free amino acids and soluble proteins was promoted in NH₄ ⁺ grown roots than NO₃ ^? grown roots at both harvesting time, whereas higher accumulation of soluble sugars was observed only at 7 days of salt stress exposure. Unlike the NH₄ ⁺ grown plants, the gas exchanges of the NO₃ ^? grown plants were reduced after 7 days of salt stress. These results suggest that external NH₄ ⁺ may limit Na⁺ accumulation in sorghum, which could contribute to improving its physiological and metabolic responses to salt stress.     

The influence of Agrobacterium rhizogenes on induction of hairy roots and ß-carboline alkaloids production in Tribulus terrestris L.

We have developed an efficient transformation system for Tribulus terrestris L., an important medicinal plant, using Agrobacterium rhizogenes strains AR15834 and GMI9534 to generate hairy roots. Hairy roots were formed directly from the cut edges of leaf explants 10–14 days after inoculation with the Agrobacterium with highest frequency transformation being 49 %, which was achieved using Agrobacterium rhizogenes AR15834 on hormone-free MS medium after 28 days inoculation. PCR analysis showed that rolB genes of Ri plasmid of A. rhizogenes were integrated and expressed into the genome of transformed hairy roots. Isolated transgenic hairy roots grew rapidly on MS medium supplemented with indole-3-butyric acid. They showed characteristics of transformed roots such as fast growth and high lateral branching in comparison with untransformed roots. Isolated control and transgenic hairy roots grown in liquid medium containing IBA were analyzed to detect ß-carboline alkaloids by High Performance Thin Layer Chromatograghy (HPTLC). Harmine content was estimated to be 1.7 μg g⁻¹ of the dried weight of transgenic hairy root cultures at the end of 50 days of culturing. The transformed roots induced by AR15834 strain, spontaneously, dedifferentiated as callus on MS medium without hormone. Optimum callus induction and shoot regeneration of transformed roots in vitro was achieved on MS medium containing 0.4 mg L⁻¹ naphthaleneacetic acid and 2 mg L⁻¹ 6-benzylaminopurine (BAP) after 50 days. The main objective of this investigation was to establish hairy roots in this plant by using A. rhizogenes to synthesize secondary products at levels comparable to the wild-type roots.     

Characterization of Phloem iron and its possible role in the regulation of fe-efficiency reactions

`Fe-efficiency reactions' are induced in the roots of dicotyledonous plants as a response to Fe deficiency. The role of phloem Fe in the regulation of these reactions was investigated. Iron travels in the phloem of Ricinus communis L. as a complex with an estimated molecular weight of 2400, as determined by gel exclusion chromatography. The complex is predominantly in the ferric form, but because of the presence of reducing compounds in the phloem sap, there must be a fast turnover in situ between ferric and ferrous (k ≈ 1 min⁻¹). Iron concentrations in R. communis phloem were determined colorimetrically or after addition of ⁵⁹Fe to the nutrient solution. The iron content of the phloem in Fe-deficient plants was lower (7 micromolar) than in Fe-sufficient plants (20 micromolar). Administration of Fe-EDTA to leaves of Phaseolus vulgaris L. increased the iron content of the roots within 2 days, and decreased proton extrusion and ferric chelate reduction. The increase in iron content of the roots was about the same as the difference between iron contents of roots grown on two iron levels with a concomitantly different expression of Fe-efficiency reactions. We conclude that the iron content of the leaves is reflected by the iron content of the phloem sap, and that the capacity of the phloem to carry iron to the roots is sufficient to influence the development of Fe-efficiency reactions. This does not preclude other ways for the shoot to influence these reactions.     

Investigations on the causes of low susceptibility of scentless mayweed to 2,4-D

White mustard (Sinapis alba L.) plants, susceptible to 2,4-D, and scentless mayweed (Tripleurospermum maritimum L.) plants, resistant to 2,4-D, markedly differ in the distribution and metabolism of 2,4-D. When 2,4-D-¹⁴C was applied onto the leaf, radioactivity was found in mustard after 8 days in the entire plant, whereas in scentless mayweed radioactivity occurred mainly in the leaf onto which it was applied and in another one or two leaves, with the roots showing only traces of radioactivity. The two plant species differed both in the character and in the number and amount of metabolites detected in aqueous and ether fractions. The metabolite with R_f 0.63 -0.65, soluble in ethylether (39% of the total radioactivity applied), and that with R_f 0.35 -0.36 (23% of the total radioactivity) were present exclusively in the roots of scentless mayweed plants. The R_f values established indicate that the former metabolite may be a conjugate with the amino acids alanine and valine and the latter a hydroxylated derivate of 2,4-D. In the shoots of both plants, 2,4-D-¹⁴C was metabolized to identical metabolites with R_f 0.59 -0.61, which occurred in mustard plants only in trace amounts. Free 2,4-D-¹⁴C occurred in the shoots of both plants and in considerable amounts also in mustard roots; it could not be demonstrated in the roots of scentless mayweed. The two species did not differ in the uptake rate or in the amount of absorbed 2,4-D-¹⁴C.     

Transfer of nitrogen from damaged roots of white clover (Trifolium repens L.) to closely associated roots of intact perennial ryegrass (Lolium perenne L)

An experiment is described in which the magnitude of N transferred from damaged white clover roots to perennial ryegrass was determined, using ¹⁵N labelling of the grass plant. There was no effect on the growth and N-fixation of the clover plants after removing part of the root system. The ¹⁵N data suggested that N had been acquired by all grass plants, even in plants grown alone with no further N supplied after labelling. However, after quantifying the mobile and stored N pools of the grass plants it was evident that significant transfer of N from clover to grass only took place from damaged clover roots. Dilution of the atom% ¹⁵N in the roots of the grass plants grown alone, and in association with undamaged clover roots, was explained by remobilisation of N within the plant.     

The use of C14O2 canopy techniques for measuring carbon transfer through the plant-soil system

Summary Methods for labelling growing plants by exposing them to C¹⁴O₂ under a cellulose acetate-butyrate canopy have been developed for laboratory and field use. The length of labelling ranged from 2 to 33 days and the C¹⁴O₂ content of the atmosphere was automatically controlled. This made it possible to measure carbon assimilation by the plants, transfer of photosynthates beneath ground and respiration of the roots.In the laboratory, root respiration of wheat plants was measured by separating the above and beneath ground plant parts using a RTV rubber partition. Half to two thirds of the assimilated carbon was found above ground, 15 to 25 per cent in the roots and shoot bases below the partition and 17 to 25 per cent was lost by underground respiration. The variability of these proportions was related to the stage of maturity of the plants.On native grassland, the relative above and beneath ground productivity was 50 per cent. The time required for the photosynthates to reach the roots at various depths ranged from 1 to 5 days and the amount of material deposited in the roots changed with time and soil moisture content. The use of tubes inserted at various depths beneath the canopy permitted sampling of soil air for C¹⁴ and CO₂ measurements. The soil C¹⁴O₂ flux indicated that root respiration during 8 days accounted for 24 per cent of the labelled carbon translocated to the roots after a two days labelling period.     

Root growth of wetland plants with different root types

A wastewater culture system was designed to study the root growth of eight species of wetland plants with two different root types. The system included a plastic barrel for holding the wastewater and a foam plate for holding the plant. The results indicated that the root growth of the plants with fibril roots was faster than that of the plants with rhizomatic roots. The species with fibril roots had higher root number (1349 per plant) than species with rhizomatic roots (549 per plant) after ten weeks of cultivation. The average root biomass of plants with fibril roots was 11.3 g per plant, whereas that of plants with rhizomatic roots was 7.4 g per plant. Fine root biomass of diameter ≤ 1 mm constituted 51.9% of the total root biomass in plants with fibril roots, whereas it accounted for only 25.1% in plants with rhizomatic roots. The root surface area of the plants with fibril roots (6933 cm² per plant) was markedly larger than that of the species with rhizomatic roots (1897 cm² per plant). The species with rhizomatic roots showed a longer root lifespan (46.6 days) than those with fibril roots (34.8 days).     

Effects of exogenous cytokinins on flowering of the short-day plantChenopodium rubrum L.

Kinetin at a concentration from 3.10^-6 M to 1.10^-3 M was applied to the plumule ofChenopodium rubrum plants during photoperiodic induction. Different levels of induction were compared (one and three short days). The higher concentrations of kinetin applied to induced plants inhibited flower formation. The rate of leaf initiation was increased under these treatments. Lower concentrations of kinetin (from 3.10^-6 M to 1.10^-5 M) usually promoted lateral bud formation and flowering. The step-wise application of kinetin revealed that the inhibitory effect on flowering had been restricted to the inductive period. The effects of kinetin, benzyladenine and trans-zeatin were compared in plants partially induced by two short days. High concentrations always inhibited flowering. Benzyladenine was the most effective in this respect. Root removal diminished the inhibitory effects of cytokinins on flowering as was stated with benzyladenine. It is assumed that endogenous cytokinins play a role in the regulation of organogenetic activity of the stem apical meristem. Depending on the photoperiodic conditions, they presumably exert their activity by maintaining the vegetative functions of the apex.     

The uptake and distribution of 15N2 into the various organs of Typha latifolia L.

Direct evidence of heterotrophic dinitrogen fixation associated with the emergent aquatic angiosperm, Typha latifolia L., was obtained through the exposure of actively growing plants to ¹⁵N₂ gas for 7 days in a gas-tight exposure vessel. Highest enrichments of ¹⁵N were found in roots/rhizomes and leaf bases. Slight enrichments were also found in the leaves due to translocation from the roots, rhizomes and leaf bases. Total fixed ¹⁵N values were 71.8 μg for the plant and 49.1 μg for the soil.Plants growing in silica sand, which received a nutrient solution containing combined nitrogen, exhibited higher enrichments and fixed 86% more ¹⁵N after exposure to ¹⁵N₂ gas than plants which received a nutrient solution lacking combined nitrogen. It is hypothesized that the concentration of combined nitrogen added was insufficient to repress nitrogen fixation and resulted in an increase in nitrogen fixation by associated microorganisms.Propane was used to trace the loss and movement of gases from the ¹⁵N₂ vessel and between the upper leaf chamber and the lower root chamber. Gas was rapidly exchanged between the upper and lower chambers through the leaves and roots of T. latifolia. Further investigations showed that propane moved at a rate of 1223 μmol day⁻¹ from the leaves to the roots and 2652 μmol day⁻¹ from the roots to the leaves. These data demonstrated that gases diffuse rapidly through the plant body of T. latifolia.     

Heterotrophic n(2) fixation and distribution of newly fixed nitrogen in a rice-flooded soil system

Rice (Oryza sativa L.) plants growing in pots of flooded soil were exposed to a ¹⁵N₂-enriched atmosphere for 3 to 13 days in a gas-tight chamber. The floodwater and soil surface were shaded with a black cloth to reduce the activity of phototrophic N₂-fixing micro-organisms. The highest ¹⁵N enrichments were consistently observed in the roots, although the total quantity of ¹⁵N incorporated into the soil was much greater. The rate of ¹⁵N incorporation into roots was much higher at the heading than at the tillering stage of growth. Definite enrichments were also found in the basal node and in the lower outer leaf sheath fractions after 3 days of exposure at the heading stage. Thirteen days was the shortest time period in which definite ¹⁵N enrichment was observed in the leaves and panicle. When plants were exposed to ¹⁵N₂ for 13 days just before heading and then allowed to mature in a normal atmosphere, 11.3% of the total ¹⁵N in the system was found in the panicles, 2.3% in the roots, and 80.7% in the subsurface soil. These results provide direct evidence of heterotrophic N₂ fixation associated with rice roots and the flooded soil and demonstrate that part of the newly fixed N is available to the plant.     

Effect of Nitrate and Ammonium Nutrition of Nonnodulated Phaseolus vulgaris L. on Phosphoenolpyruvate Carboxylase and Pyruvate Kinase Activity

Young bean plants (Phaseolus vulgaris L. var Saxa) were fed with 3.5 or 10 millimolar N in either the form of NO₃⁻ or NH₄⁺, after being grown on N-free nutrient solution for 8 days. The pH of the nutrient solutions was either 6 or 4. The cell sap pH and the extractable activities of phosphoenolpyruvate carboxylase and of pyruvate kinase from roots and primary leaves were measured over several days.

The extractable activity of phosphoenolpyruvate carboxylase (based on soluble protein) from primary leaves increased with NO₃⁻ nutrition, whereas with NH₄⁺ nutrition and on N-free nutrient solution the activity remained at a low level. Phosphoenopyruvate carboxylase activity from the roots of NH₄⁺-fed plants at pH 4 was finally somewhat higher than from the roots of plants grown on NO₃⁻ at the same pH. There was no difference in activity from the root between the N treatments when pH in the nutrient solutions was 6. The extractable activity of pyruvate kinase from roots and primary leaves seemed not to be influenced by the N nutrition of the plants.

The results are discussed in relation to the physiological function of both enzymes with special regard to the postulated functions of phosphoenolpyruvate carboxylase in C3 plants as an anaplerotic enzyme and as part of a cellular pH stat.

     

Distribution of photoassimilates in the pea plant: chronology of events in non-fertilized ovaries and effects of gibberellic acid

The short-lived isotope¹¹C (t_1/2=20.4 min) has been used to study assimilate distribution in intact pea plants (Pisum sativum L.). Radiolabel was measured at the leaf fed with¹¹CO₂ (feed-leaf), at the ovary of the flower subtended by this leaf, and in shoot apex and roots of individual plants. Considerable¹¹C-radiolabel was detected in the young ovaries during the first days after anthesis. Thereafter, when the ovaries stopped growing the uptake of¹¹C rapidly decreased. At this developmental stage only apex and roots were competing for the photoassimilates. Fertilization, however, restored the strong sink activity of the ovaries. The same effect could be achieved by applying gibberellic acid to non-fertilized ovaries. About 2 h after treatment the residual¹¹C-radiolabel entering the ovary started to increase and, at about the same time, the ovary resumed growth. Feed-leaf photosynthesis, as well as export of¹¹C-radiolabel out of the leaf, was not changed by the treatment. The¹¹C experiments show the dynamic behaviour of the sinks during developmental stages from the day of anthesis until 5 d later and demonstrate that phytohormones may play an important role in regulating carbon distribution.