46 JOURNAL OF APPLIED SEED PRODUCTION, VOL. 3, 1985this was probably located in the developing seeds (Fig. 3c).The assimilate supply to seeds was not recorded separately inthis experiment but in a similar study with Poa annua approximately70% of the t4C in the inflorescence was recoveredfrom seeds (Ong and Marshall, 1975). During anthesis theterminal internode was the major sink for assimilate exportedfrom the ryegrass inflorescence and this suggests that theremay be competition for assimilate between newly developingseeds and the elongating internode. Assimilate exported fromthe flag leaf and lower leaves was utilized almost entirely bythe stem (Fig. 3 a, b) and thus in contrast to most other workon source-sink relationships in grasses and cereals (Ong andMarshall, 1975; Wardlaw, 1975) these leaves did not make asignificant contribution to the growth and development ofseeds. They did, however, support the development of theinflorescence in the pre-anthesis period.Although these results suggest that the photosynthetic604020(a)inflorescence appearanceanthesis1----1seed fillingseed ripeningactivity of the inflorescence is wholly adequate to supportseed growth, observations in another season revealed that20% of the t4C in plants four weeks after supplying t4CQ 2tothe entire tiller at anthesis was recovered from the seeds (Fig.4). Thus the current assimilation of the inflorescence may besupplemented by the mobilization of material previouslyassimilated by the leaves and temporarily stored in the lowerinternodes in the post-anthesis period as described for wheat(Austin et al.,l977).10080I 601l~ 40[u::';(a)inflorescence appearancean thesisseed filling• 0 ••....·::.·seed ripening040(b)0(b) 1002040602080060(c)40202000 10 2030 40Days from inflorescence appearance50 60Figure 3. The percentage distribution of 14C-labelled assimilateafter 24 hours from (a) flag leaf, (b) penultimate (PL) or basal(BL) leaf, and (c) inflorescence, at different stages of reproductivedevelopment ( + S.E. ). IS] ,inflorescence; ~ , leavesand upper internode; D , rest of stern; IIIII , other tillers; ill] ,root. (From Colvill and Marshall, 1984).04060Days from inflorescence appearanceFigure 4, The redistribution of 14C-labelled assimilate with timein L. perenne. Main tiller supplied with 14{;0 2 at (a) flag leafemergence, or (b) anthesis. EJ , inflorescence; 0 , upperinternode; 0 , rest of stern and leaves; § , seeds;• , other tillers; IIIII1 , root. (From Colvill & Marshall,1984).
JOURNAL OF APPLIED SEED PRODUCTION, VOL. 3, 1985 47A recently reported field study on I4C-assimilate distriutionin L. perenne during seed filling by Clemence andHebblethwaite (1984) shows an interesting additionalsource-sink link compared with the previously describedstudy of Ong et al. ( 1978a) and Colvill and Marshall (1984).The latter was conducted on a sward supplied with a relativelylow input of nitrogen fertilizer to minimize lodging andobservations were made in an unusually dry season, whereasHebblethwaite and co-workers studied crops well suppliedwith nitrogen. The higher input of nitrogen fertilizer stimulatestillering as well as causing early and extensive lodging(Hebblethwaite and Ivins, 1977; Hebblethwaite eta!., 1980)and so in this situation young tillers which rely on importedassimilate for their development (Colvill and Marshall, 1981)will form a significant sink for assimilate, a demand that canonly be met by established flowering tillers. Correspondingly,Clemence and Hebblethwaite (1984) found that I4C-assimilatewas readily supplied to developing tillers by theinflorescence and flag leaf during seed filling whereas Ong etal. (1978b) and Colvill and Marshall (1984) recorded only atrace of assimilate moving out of a flowering tiller to subtendingtillers at this time. However, inPoa annua, Ong andMarshall (1975) also found that young tillers were a majorsink for assimilate from the inflorescence. In contrast, Colvilland Marshall (1984) found that developing tillers were supportedby the mobilization of reserve materials rather than bycurrent assimilate as one sixth of the I4C in a plant four weeksafter supplying I4C0 2 to the shoot at anthesis was localized innew tillers (Fig. 4). In a lodged rye grass crop the flag leaf andother leaves thus supply assimilate to two major sinks, developingtillers and the stem, and as a result there is onlyminor support to growing seeds, but even so both Clemenceand Hebblethwaite (1984) and Hampton and Hebblethwaite( 1985b) recorded a relatively greater allocation of assimilatefrom the flag leaf to the inflorescence than that observed byColvill and Marshall (1984).It is evident from these I4C-studies that the inflorescence isthe principal assimilatory organ of the flowering tiller andsupplies the majority of the assimilate utilized by developingseeds. A significant proportion of current assimilate is exportedfrom the inflorescence and in view of this Colvill andMarshall (1984) concluded that the production of assimilatewas far greater than that needed for seed growth. This view issupported by experiments where lodging is prevented eitherby mechanically supporting the crop (Burbidge et a!., 1978)or chemically (Wright and Hebblethwaite, 1979; Hamptonand Hebblethwaite, 1985b) where the growth and developmentof many more seeds per inflorescence can be supported,but this response may in part reflect an improvement in thewithin-crop environment. However, the setting of seeds andtheir successful early growth is a far more critical developmentalstage than their overall growth to maturity and thus ifthere is some degree of competition between vegetative andreproductive sinks for assimilate (mainly from the inflorescence)then this could perhaps influence the abortion ofdeveloping seeds.Effects of Growth Regulators and FungicidesAny experimental treatment that modifies source-sinkrelations to the advantage of reproductive growth and development,for example the application of growth regulatorsthat reduce stem development and thereby reduce lodging,might be expected to enhance floret and seed development.Whilst the results of experiments with growth regulators suchas Ancymidol and Paclobutrazol indeed show large increasesin seed yield, mainly associated with an increase in seednumber per spikelet, the mechanism underlying this improvementis not clear (Hebblethwaite eta!., 1982; Hamptonand Hebblethwaite, 1985b).In particular, three aspects need to be resolved. Firstly,whether the number of fertile florets per spikelet at an thesis isinfluenced by the application of growth regulators: someobservations suggest that this is unaltered (Wright andHebblethwaite, 1979) but others show a significant increase(Hampton and Hebblethwaite, 1985b). Secondly, whetherthe reduction in lodging of the crop associated with theapplication of chemicals provides a more favorable environmentfor the production, release and dispersal of pollen,and so correspondingly results in an increase in the proportionof florets that become fertilized and set seed (Griffiths eta!., 1980; Hill, 1980). Thirdly, whether the loss of developingseeds by abortion is reduced by the altered growth patternof the crop following the application of growth regulators.This has frequently been claimed but is not supported bydetailed observations on seed number per spikelet with time.However, in a recent publication Hampton and Hebblethwaite( 1985b) provide some evidence that Paclobutrazol applicationat the time of spikelet initiation significantly reduced theabortion of seeds. This may reflect a direct effect of thegrowth regulator in suppressing the growth of vegetativetillers that compete with developing seeds for assimilate, orindirectly reflect improved light relations within the cropassociated with reduced lodging, i.e. tillers may, by virtue ofenhanced photosynthetic capacity, make less demand forassimilate from other parts of the plant.In general it can be concluded that competition for assimilatebetween reproductive and vegetative sinks especiallyduring anthesis and the early period of seed growth is importantin regulating seed yield. This view is supported by theresults of recent experiments in which fungicides have beenapplied to lodged crops of ryegrass; the crop remains lodgedand seed production per spikelet is significantly increased(Hampton and Hebblethwaite, 1984). This increase in seedyield was brought about in the virtual absence of pathogenicfungi and was related to an increase in the leaf area durationof the canopy resulting from delayed leaf senescence. Astreatment with fungicide did not influence floret number perspikelet at anthesis and did not alter the structure of the cropcanopy, the improvement in seed production per inflorescencemust largely be due to a reduction in seed abortion. Itcan, therefore, be suggested that treatment with fungicidemaintained the photosynthetic activity of tillers and resultedin an improved supply of assimilate to developing seeds at atime when they are normally prone to abort. In addition it ispossible that fungicides also exert a direct hormonal effect ondeveloping seeds via a cytokinin-like action. This wouldstimulate the sink activity of developing seeds and sustaintheir import of assimilate and other substrates, thereby re-
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