Free amino Acid content and metabolic activities of setting and aborting soybean ovaries

Fruits of soybean (Glycine max [L.] Merr.) that are destined to abscise shortly after anthesis grow more slowly than fruits that will be retained. In this work, amino acid composition, protein metabolism, and nucleic acid metabolism were studied in setting and abscising soybean ovaries from anthesis to 6 days after anthesis to provide additional evidence of chemical processes associated with abscission. Principal free amino acids were asparagine, aspartic acid, glutamic acid, serine, and glutamine. Percent aspartate and glutamate declined as the ovaries grew, with aspartate declining more in abscising and glutamate more in setting ovaries. Percent glutamate was positively correlated to percent abscission throughout the period. Proline, serine, and leucine were positively correlated to abscission from 0 to 2 days after anthesis, whereas significant negative correlations were observed at these ages for ethanolamine and arginine. ⁷⁵Se fed as selenate and ¹⁴C fed as sucrose, glycine, and alanine were readily incorporated into soluble and insoluble proteins in a 24-hour in vitro incubation. Radioactivity of total proteins, expressed on a perovary basis, was negatively correlated with percent abscission and positively correlated with ovary weight. [¹⁴C]Glutamine and serine followed the opposite pattern, with greater protein labeling in abscising than in setting ovaries. When data were expressed as disintegrations per minute per milligram ovary fresh weight, protein labeling from alanine was seen to be significantly greater in abscising ovaries at anthesis and throughout the sampling period. Nucleic acid labeling from uridine was highly correlated to ovary weight; labeling from thymidine was greater in setting than abscising ovaries at anthesis and in abscising ovaries at later stages of development. It is concluded that abscising ovaries can continue amino acid metabolism almost up to the date of separation from the raceme, and that the involvement of alanine, glutamine, aspartate, glutamate, and other amino acids in soybean flower abortion deserves further study.     

Biochemical characterization of soybean ovary growth from anthesis to abscission of aborting ovaries

Soybean (Glycine max [L.] Merr.) ovary growth was measured from anthesis to 6 days after anthesis (DAA) to establish a timetable of biochemical events that might be useful in identifying processes that initiate abscission. Two procedures were developed to provide samples with either high or low percent pod set for `IX93-100,' a semideterminate line having long racemes. Characteristics measured were fresh and dry weight, soluble and insoluble protein, soluble carbohydrate, starch, RNA, and DNA. Setting ovaries grew more rapidly than abscising ovaries. Since there was a daily increase in ovary weight in both groups, all measured characteristics showed daily increases when expressed on perovary basis. Statistically significant differences between groups were detected between 2 and 5 DAA for most characteristics. When chemical composition was expressed on concentration basis, starch level was significantly higher in setting ovaries at 5 and 6 DAA. Regression analysis showed that these deviations between setting and abscising samples started between anthesis and 1 DAA. We conclude that processes leading to eventual shedding of fertilized ovaries (called flower abortion in soybeans) commence soon after anthesis of the shed flower, and that setting and abscising ovaries do not differ in protein, soluble carbohydrate, starch, or nucleic acid content when abscission processes begin.     

Light and Shade Effects on Abscission and C-Photoassimilate Partitioning among Reproductive Structures in Soybean

Field experiments were conducted in 1981 and 1982 to study the effects of low-irradiance supplemental light on soybean (Glycine max [L.] Merr. cv Evans) flower and pod abscission. Cool-white and red fluorescent lights illuminated the lower part of the soybean canopy during daylight hours for 3 weeks late in flowering. At the same time, flowers and young pods on half the plants were shaded with aluminum foil. Flowers were tagged at anthesis and monitored through abscission or pod maturity.

Responses to red and white lights were similar. Supplemental light tended to reduce abscission and increase seed weight per node compared to natural light. Shading flowers and pods increased abscission and reduced seed weight per node. Number of flowers produced per node, individual seed weight, and seeds per pod were not affected by light or shade treatments.

Further studies examined the effects of shading reproductive structures on their capacity to accumulate ¹⁴C-photoassimilates. Individual leaves were pulse labeled with ¹⁴CO₂ 1, 2, and 4 weeks post anthesis. Flowers and pods in the axil of the labeled leaf were covered with aluminum foil 0, 24, 72, and 120 hours before pulsing.

Shading flowers and pods resulted in a 30% reduction in the relative amount of radiolabel accumulated from the source leaf. The reduction in ¹⁴C accumulation due to shading was evident regardless of the length of the shading period and was most pronounced when the shades were applied early in reproductive development. We conclude that light perceived by soybean flowers and young pods has a role in regulating both their abscission and their capacity to accumulate photoassimilates.

     

C]Ethylene Metabolism during Leaf Abscission in Cotton

Changes in ¹⁴C₂H₄ metabolism in the abscission zone were monitored during cotton (cv. Deltapine 16) leaf abscission. Rates of ¹⁴C₂H₄ oxidation to ¹⁴CO₂ and tissue incorporation in abscission zone segments cut from the second true leaf of nonabscising leaves of intact plants were similar (about 200 disintegrations per minute per 0.1 gram dry weight per 5.5 hours) and relatively constant over a 5-day period. Deblading to induce abscission caused a dramatic rise in ¹⁴C₂H₄ oxidation, but tissue incorporation was not markedly affected. This rise occurred well before abscission, reaching a peak of 1,375 disintegrations per minute per 0.1 gram dry weight per 5.5 hours 2 days after deblading when abscission was 40%. The rate then gradually declined, but on day 5 when abscission reached completion, it was still nearly three times higher than in segments from nonabscising leaves. Application of 0.1 millimolar abscisic acid in lanolin to the debladed petiole ends increased the per cent abscission slightly and initially stimulated ¹⁴C₂H₄ oxidation. In contrast, naphthaleneacetic acid applied in a similar manner delayed and markedly inhibited both abscission and ¹⁴C₂H₄ oxidation.     

Changes in Abscisic Acid and Indoleacetic Acid before and after Anthesis Relative to Changes in Abscission Rates of Cotton Fruiting Forms

Cotton (Gossypium hirsutum L.) fruiting forms exhibit pronounced changes, with age, in their probability of abscission. Large floral buds rarely abscise, but after anthesis the young fruits (bolls) have a high probability of abscising. Abscission rate reaches a peak about 5 to 6 days after anthesis and then gradually decreases. An experiment was conducted to try to determine the reason for the rapid and pronounced increase in probability of abscission just after anthesis. Cotton was grown in the field and fruiting forms of various ages from 9 days before to 9 days after anthesis were all harvested the same day and subsequently analyzed for ABA and IAA. The concentration of ABA decreased slightly at anthesis and increased gradually thereafter. In contrast, the concentration of IAA was high before anthesis and then decreased at anthesis to about one-fifth the previous concentration. IAA remained low for at least 4 days after anthesis and then increased rapidly between 7 and 9 days after anthesis. The high concentration of IAA in floral buds before anthesis is probably a major factor in their resistance to abscission. Likewise, the low concentration of IAA at anthesis and for about 4 days thereafter may promote fruit abscission during the young boll stage.     

Fruit Abscission in Cowpea, Vigna unguiculata (L.) Walp: I. DISTRIBUTION OF 14C-ASSIMILATES IN THE INFLORESCENCE, AND COMPARATIVE GROWTH OF OVARIES FROM PERSISTING AND ABSCISING OPEN FLOWERS

One of the hypotheses proposed for the abscission of flowersand fruits from the cowpea was that the oldest fruits at raceme1 on the peduncle monopolize the mobile nutrients availableto the whole inflorescence. Consequently, the distal young fruitsand flowers starve, abort, and are shed. This hypothesis wastested through autoradiographic studies of the distributionof ¹⁴C-assimilates on the cowpea inflorescence and the comparativegrowth of ovaries from persisting and abscising open flowersand young fruits. ¹⁴C-asaimilates were distributed to the fruits at raceme 1 andto flower buds, open flowers, and young fruits distal to raceme1. Therefore, the hypothesis that the fruits at raceme 1 monopolizethe mobile assimilates cannot be maintained. At the time of opening the persisting flowers of raceme 1 hadheavier ovaries than the aborting flowers of raceme 3 althoughboth types of flowers were visually alike. During the initial12 h following opening, the difference in ovary size was maintainedand often accentuated. The ovary of the persisting flower increasedin both fresh and dry weights. The growth of the ovary of theaborting flower was variable; the fresh weight often decreased,although it also increased in some experiments; the dry weightremained unchanged or decreased or increased remarkably. The availability of nutrients was considered to be of secondaryimportance in the chain of events leading to fruit abscission.Other internal factors which may control vital processes relatedto embryo development appear to be of primary, underlying importance.     

The roles of abscisic acid and ethylene in the abscission and senescence of cocoa flowers

Cocoa flowers have a limited period of longevity; more than 90% of unpollinated flowers abscised within 32 h after anthesis. Abscisic acid (ABA) levels increased significantly prior to abscission. By 21 h after anthesis, ABA levels had increased almost 10-fold, and by 32 h flowers had 20-fold higher levels of ABA than at anthesis. Fluridone completely inhibited both the increase in ABA, the formation of an abscission zone, and the abscission and senescence of flowers. In contrast, ethylene production increased only slightly 21 h after anthesis and was only 2-fold higher after 32 h. Aminoethoxyvinylglycine (AVG) delayed but did not prevent abscission. In cocoa flowers, ABA is the primary regulator of abscission; ethylene accelerates abscission but only in the presence of ABA. Naphthalene acetic acid (NAA) treatment of flowers at anthesis prevented abscission zone formation and flower abscission, but did not induce fruit set. All parts of the NAA-treated flower except the pedicel senesced after 6 days. NAA+AVG treatment only delayed, whereas fluridone treatment completely prevented flower senescence.     

Hormonal Regulation of Pedicel Abscission in Begonia Flower Buds

In order to analyse the hormonal regulation of flower bud shedding in Begonia, levels of indoleacetic acid (IAA), abscisic acid (ABA) and ethylene were determined in buds and pedicels. The translocation and metabolism of ¹⁴C-labeled IAA in pedicel segments were also studied. In a monoecious Begonia fuchsioides hybrid, abscising male flower buds contain about 1% of the IAA present in non-abscising female flowers. In a male Begonia davisii hybrid, the seasonal variation in bud drop coincides with changes in the IAA content of the buds, while also the release of IAA from the bud to the pedicel is hampered. Abscission zones of these pedicels always contain abscission promoting ethylene concentrations. The tissue is prevented from responding with abscission by IAA from the flower buds. The buds also contain ABA but without influencing abscission considerably. Pretreatment with ethylene or ABA does not affect IAA transport in pedicel segments. The rate of this transport is 4–6 mm × h^–1:; the capacity increases with the transverse area. In young segments, IAA is decarboxylated and also otherwise metabolized.     

Fruit growth and abscission in Macadamia imtegrifolia

Growth in diameter of fruits (follicles) of five cultivate of Macadamia integrifolia Maiden and Betche followed a single sigmoidal pattern. A rapid increase in fruit diameter began 2 to 3 weeks after anithesis and continued until weeks 14 to 16. No further increase in size was observed after 16 weeks. Growth in fresh weight of the two cnltivars observed also followed a single sigmoidal pattern. A rapid increase in fresh weight began 5 to 6 weeks after anthesis and continued until weeks 15 to 18. No further increase in fresh weight was evident after 18 weeks. Abscission of flowers and fruits occurred continuously from anthesis through fruit maturity and could be separated into three periods. The first period of abscission occurred in the first 2 weeks after anthesis and commonly involved over 90% of the flowers. The second period of abscission occurred from 3 to 8 weeks after anthesis and commonly involved over 80% of the initial set of small fruit. This period of abscission occurred prior to and concurrent with the rapid increase in growth. The third period of abscission was gradual and occurred from 9 weeks after anthesis until maturity at 28 to 30 weeks. Measurement of ethylene evolution by fruits showed a peak during weeks 3 and 4. This was prior to the peak in relative abscission rate at week 5. The fruit removal force increased from anthesis through week 12 and decreased from week 24 through maturity.     

Metabolic depression and Na+/K+ gradients in the aestivating Australian goldfields frog,Neobatrachus wilsmorei

During aestivation the metabolic rate of the Australian goldfields frog Neobatrachus wilsmorei was reduced by 80% from its standard metabolic rate. The in vitro rate of oxygen consumption of isolated muscle and skin from aestivating frogs was up to 50% lower than that of the non-aestivating frogs. This in vitro rate of oxygen consumption was maintained for 6–12 h, indicating an intrinsic metabolic depression of tissues during aestivation. Frogs became dehydrated during aestivation. Muscle, skin and liver also became dehydrated during aestivation, but brain and kidney did not. Na⁺ and K⁺ contents and extracellular space measurement for muscle indicated that ion gradients were maintained across the muscle cell membrane during aestivation. Increases in plasma concentrations of Na⁺ and K⁺ were matched with similar increases in muscle intracellular ion concentrations. Extracellular space measurements were unsuccessful in the other tissues, but K⁺ content in all tissues (per dry weight) was maintained during aestivation, and the concentration of plasma K⁺ did not increase above that which can be accounted for by dehydration, indicating that K⁺ gradients were maintained.Abbreviations bm body mass - DPM disintegrations per minute - dw dry weight - MR metabolic rate - vO₂ rate of oxygen consumption - ww wet weight     

Auxins,ABA and gibberellin-like activity in abscising and non-abscising flowers and pods of Vicia faba L.

Abscission probability varies among floral positions within inflorescences of Vicia faba L. Flowers from proximal positions have a greater chance to develop into mature pods than flowers from more distal positions which normally abscise either as older flowers or as young pods. In three field experiments with the indeterminate single stem variety Herz-Freya, changes in the contents of extractable auxins, abscisic acid (ABA) and gibberellins in flowers and pods during their development, and their possible influence on abscission were investigated.Inflorescences at different positions along the stem were divided into the two proximal and the remaining fruits. The content of all three hormones was at a low level during flower development, increased greatly in parallel with dry matter accumulation in the young pods, and then decreased to maturity. The first hormone to increase in the fruits was auxin and this took place when abscission from the distal positions began. ABA and gibberellins at this time were still at a low level. This ontogenic course of hormone production was very similar in fruits of both positions within an inflorescence, but in flowers and young pods from proximal positions, auxin content in most inflorescences was greater than in those from the abscising distal positions. No such positional differences were observed with ABA and gibberellins. Decapitation of the plants reduced flower and pod drop from the remaining reproductive nodes. Although decapitation resulted in less abscission among distal flowers and young pods from these nodes, it did not affect the ontogenic course of auxin and ABA production in these fruits.     

Role of Ethylene in Senescence of Petals--Morphological and Taxonomical Relationships

Petal senescence in mature flowers was studied in 93 speciesfrom 22 families. The initial symptom of senescence was eitherwilting or abscission, but in some species the time span betweenwilting and abscission was very short. There was no apparent relationship between corolla form (choripetalousor sympetalous), ovary position (inferior or superior with respectto the corolla) and type of senescence (initial wilting or initialabscission). In monocots no initial abscission was found, whilein dicots the difference between the wilting type and the abscissiontype was generally at the family level. With respect to petalsenescence, sensitivity to exogenous ethylene (C₂H₄) was alsorelated to the family level. Except for a few families (all tested Campanulaceae, Caryophyllaceaeand Malvaceae, and most Orchidaceae), most of the flowers investigatedthat showed initial wilting were not sensitive to exogenousethylene, e.g. all tested Compositae, Iridaceae, and Liliaceae.Most of the flowers showing initial abscission were sensitiveto exogenous ethylene (Geraniaceae, Labiatae, Ranunculaceae,Rosaceae, Scrophulariaceae). Experiments with silver thiosulphate (STS) confirmed the effectsof exogenous ethylene, both in flowers showing initial wiltingand in flowers showing initial abscission. The data indicate,therefore, that ethylene is involved in the natural senescenceof only a minority of the wilting type of flowers and in a majority(if not all) of the abscising type of flowers. Key words: Abscission, ethylene, senescence, silver thiosulphate     

Life-cycles of three populations of Potamogeton pectinatus L. at different degrees of wave exposure in the Askö area,Northern Baltic proper

Potamogeton pectinatus L. populations were studied at three stations over an annual cycle.The maximum standing crop of 17.5 g dry weight (wt.) m⁻² was recorded a the most sheltered station, decreasing slightly at the intermediate station to 15.8 g dry wt.m⁻², and being only 4.8 g dry wt.m⁻² at the most exposed station.Flowering and seed production occurred at all three stations. The maximum proportion of dry weight allocated to reproduction was higher at the more exposed stations (4%), while at the sheltered station only 1% was allocated to reproductive parts. The number of overwintering tubers produced per m² decreased from 115 per m² to 45 per m² with lowered exposure. Conversely, the number of flowers per inflorescence and flowers per m² was highest at the most sheltered station, decreasing from 9 to 6 and 945 to 672 per m², respectively, with increasing exposure. No definite trend could be found for seed production, which was highly variable and dependent on a successful period of fertilization during calm weather. At the sheltered locality more resources were allocated to vegetative shoots and rhizomes which were also able to overwinter.The resource allocation of Potamogeton pectinatus seems to be strongly influenced by environmental factors. This is discussed in relation to current reproductive strategy concepts.     

Effect of light and ontogenetic stage on sink strength in bean leaves

Light (about 3,000 foot-candles) neither increased nor decreased the sink strength of young, rapidly expanding leaves of Phaseolus vulgaris L. cv. Black Valentine, as measured by the comparative rates of import of ¹⁴C-labeled photosynthates by sink leaves in the light versus dark in short term experiments. Although irradiated sink leaves accumulated more ¹⁴C activity, the difference was fully accounted for by photosynthetic reabsorption of respiratory CO₂ derived from substrates translocated to the sink leaves.

Maximum sink strength was attained when the sink leaf reached 7 to 8 cm² in area (9 to 10% of its fully expanded size). Thereafter sink strength declined rapidly and asymptotically to a near zero value at about 45% final area. During this period, however, the rapid decline in translocation was offset by a rapid rise in the photosynthetic rate of the sink leaf, maintaining a near constant relative rate of dry weight increase until the sink leaf had expanded to about 17% of its final area. Although the increasing photosynthetic capacity was associated with a decreasing import capacity, suggesting that the rate of translocation to the sink leaf was controlled by the developing capacity of the sink leaf for photosynthesis, it was not possible to vary the total (true) translocation rate to the sink leaf by varying the photosynthetic rate of the sink leaf in short term light-dark experiments. Despite a high ratio of source to sink in these experiments, no evidence accrued that translocation into young bean leaves was ever sink-limited.

     

Influence of phloem transport on flower abscission in Hibiscus rosa-sinensis

Summary Flowering cultivars of Hibiscus rosa-sinensis L. were either cross-pollinated or self-pollinated. Fruit set was observed on 52% of the cross-fertilized flowers, while only 4.6% of the self-fertilized flowers were not abscised. Once during fruit and seed growth, the subtending leaf was exposed to ¹⁴CO₂, and translocation of labelled photoassimilate was recorded by macro- and microautoradiography. Phloem transport into the raphe occurred in both fruits with fertilized and fruits with non-fertilized ovules. Since empty ovules showed some sink strength, it is assumed that growth of vegetative seed-tissue signalizes the retardation of completion of the abscission process. During fruit growth a considerable amount of starch is deposited in the distal layer of the abscission zone. Part of this starch is consumed during growth of cross-fertilized fruits.     

Site of clomazone action in tolerant-soybean and susceptible-cotton photomixotrophic cell suspension cultures

Studies were conducted to determine the herbicidal site of clomazone action in tolerant-soybean (Glycine max [L.] Merr. cv Corsoy) (SB-M) and susceptible-cotton (Gossypium hirsutum [L.] cv Stoneville 825) (COT-M) photomixotrophic cell suspension cultures. Although a 10 micromolar clomazone treatment did not significantly reduce the terpene or mixed terpenoid content (microgram per gram fresh weight) of the SB-M cell line, there was over a 70% reduction in the chlorophyll (Chl), carotenoid (CAR), and plastoquinone (PQ) content of the COT-M cell line. The tocopherol (TOC) content was reduced only 35.6%. Reductions in the levels of Chl, CAR, TOC, and PQ indicate that the site of clomazone action in COT-M cells is prior to geranylgeranyl pyrophosphate (GGPP). The clomazone treatment did not significantly reduce the flow of [¹⁴C]mevalonate ([¹⁴C]MEV) (nanocuries per gram fresh weight) into CAR and the three mixed terpenoid compounds of SB-M cells. Conversely, [¹⁴C]MEV incorporation into CAR and the terpene moieties of Chl, PQ, and TOC in COT-M cells was reduced at least 73%, indicating that the site of clomazone action must be after MEV. Sequestration of clomazone away from the chloroplast cannot account for soybean tolerance to clomazone since chloroplasts isolated from both cell lines incubated with [¹⁴C]clomazone contained a similar amount of radioactivity (disintegrations per minute per microgram of Chl). The possible site(s) of clomazone inhibition include mevalonate kinase, phosphomevalonate kinase, pyrophosphomevalonate decarboxylase, isopentenyl pyrophosphate isomerase, and/or a prenyl transferase.     

Effects of crop thinning and reduced grain numbers per ear on grain size in two winter wheat varieties given different amounts of nitrogen

To study the importance for final grain size in wheat (Triticum aestivum, L.) of assimilate supply and the storage capacity of the grain, two field experiments were done. In 1976 nitrogen was applied in the range from none to 180 kg ha^-1, part of the crop was thinned, and the top halves of some ears of the short variety Hobbit and of the tall variety Maris Huntsman were removed soon after anthesis. In 1977 ears of Maris Huntsman were halved 5 days after anthesis or at 30 days after anthesis when grain volume was maximum. Thinning the crop from 360 to 180 ear-bearing shoots m^-2 30 days before anthesis increased the number of grains per ear, except in the absence of nitrogen fertiliser, but did not increase grain size, grain dry weight per ear or total dry weight per culm. Removing the upper half of ears of Hobbit 5 days after anthesis increased dry weight per grain, but when this treatment was applied to Maris Huntsman either 5 days after anthesis in 1976 and 1977, or when grain volume was maximal in 1977, the grains failed to increase in dry weight. Non-grain dry weight of both varieties was increased by halving the ear. In both varieties the maximum volume of grains in halved ears was larger than in intact ears. Grain dry weight increased relatively less than volume after halving the ear of Hobbit, and the decrease in volume up to maturity was greater in halved than intact ears of both varieties. The larger grain volume in halved ears of Maris Huntsman in 1977 was associated with more endosperm cells.     

Changes in Amide-Linked and Ester Indole-3-Acetic Acid in Cotton Fruiting Forms during Their Development

The concentration of free indoleacetic acid (IAA) is high in cotton (Gossypium hirsutum L.) fruiting forms before anthesis, but is low at and for a few days after anthesis. Amide-linked and ester IAA were measured in fruiting forms at 9, 6, and 3 days before anthesis; at anthesis; and at 2, 4, 7, and 9 days after anthesis to determine if free IAA decreased because it was converted to a conjugated form. That did not appear to be the case. While the major decrease in free IAA occurred during the 6 days before anthesis, ester IAA increased only a small amount and amide-linked IAA decreased even more than free IAA. During the 6 days before anthesis free IAA decreased from 0.62 to 0.12 micrograms per gram and amide-linked IAA decreased from 19.14 to 1.16 micrograms per gram dry weight. No evidence was found that a large amount of amide-linked IAA was converted to an insoluble form; flowers contained less than 1 microgram per gram of insoluble IAA. The free and amide-linked IAA must have been converted to other forms, perhaps by oxidation. Soluble amide-linked IAA remained low after anthesis. No ester IAA was detected 6 days before anthesis and only 0.08 microgram per gram dry weight was measured at anthesis. The concentration of ester IAA increased thereafter to 4.43 micrograms per gram at 9 days after anthesis. Therefore, amide-linked IAA was the major form of IAA in flower buds and ester IAA was the major form in young fruits (bolls). Minimum concentrations of free and total IAA occurred during the 4 days after anthesis, a stage when cotton fruiting forms are most likely to abscise. The large decreases in free and amide-linked IAA during the 6 days before anthesis may indicate a rapid turnover of IAA in flower buds. But, the decrease in free IAA was not accompanied by a comparable increase in ester or amide-linked IAA.     

An investigation into the relations between some growth parameters and yield of barley

Twenty-five field experiments on barley were done at four sites from 1970 to 1975, with an additional site in 1975, comparing five rates of nitrogen application. The crops were sampled at c. 14-day intervals from about the four-leaf stage to maturity to follow changes with time in the dry weights of the constituent plant parts and to measure fertile tiller number, grain number per ear and grain weight. Some data have been selected from this body of information and regression analysis has been used to assess the possible importance of pre- and post-anthesis growth in determining the yield of barley. Ear emergence was taken as an approximate indicator of anthesis. Yield was closely related (r=+0–96) and almost equal to the amount of dry matter accumulated before ear emergence for twenty-one of the experiments, with a regression coefficient of 0–95. In two of the remaining experiments the regression coefficient was greater, 1–14; and smaller in the other two, o-8i. Plant population is also shown to affect the regression coefficient. Despite these anomalies grain yield was more closely related to plant dry weight at ear emergence than to the increase in weight after ear emergence (r =+0–79). The increase in plant dry weight after ear emergence was frequently less than the yield of grain, suggesting that the plant can compensate for inadequate photosynthesis during grain filling. The result of this compensation is a reduction in straw dry weight. This implies that previously assimilated dry matter is transported to the grain, or that respiratory losses from the straw are not replaced by current photosynthesis. These observations suggest that yield may be limited by sink capacity rather than by photosynthesis after ear emergence. The main component of sink capacity, grain number per m² was closely related to yield (r =+0–95) and is known to be determined at or before anthesis. Grain size was shown to be related to grain number per ear (r= 4- 0–99), suggesting that grain size is also, at least partially, determined at ear emergence. Therefore, all the yield components are determined, wholly or partly before ear emergence.