Effect of the Duration of the Vegetative Phase on Shoot Growth, Development and Yield in Pearl Millet (Pennisetum americanum (L.) Leeke)

Craufurd, P. Q. and Bidinger, F. R. 1988. Effect of the durationof the vegetative phase on shoot growth, development and yieldin pearl millet (Pennisetum americanum (L.) Leeke).–J.exp. Bot. 38: 124–139 The duration of the vegetative phase (DVP) in millet, whichis the major cause of variation in the crop duration, has markedeffects on the number of productive tillers per plant and onmainshoot (MS) and tiller grain yield. Daylength extensionswere used to vary the DVP and the effect on factors affectingpanicle (tiller) number per plant and panicle yield examinedin millet hybrid 841A x J104, grown in the field at Hyderabad,India. Tiller appearance, shoot leaf appearance and leaf area,and stem and panicle growth, in both MS and primary tillers(PTs), were monitored at frequent intervals over the season.At maturity grain yield per shoot was measured The concept of thermal time was used to describe shoot development.The rates of tiller appearance and shoot leaf appearance werelinearly related to thermal time and were not affected by DVPtreatments. The duration of the growth phase from panicle initiationto flowering (GS2) and from flowering to maturity (GS3) was320 and 390 degree days (°Cd), respectively. There was nodifference in rates of leaf or tiller appearance or developmentbetween MS and PTs. Tiller appearance, tiller leaf appearanceand tiller apical development all ceased at the same time inthe later initiated PTs, approximately 550 °Cd from sowing,shortly after rapid stem growth had begun. Tillers that didnot survive were all vegetative or in the early stages of reproductivedevelopment at this time The rate of accumulation of dry matter per plant was similarin all DVP treatments, but in the longer DVP treatments a greaterproportion of the dry matter was partitioned to the MS. Mainshootstem and panicle growth rates were increased by a longer DVP,as was grain yield on the MS, and these were related to increasedMS leaf area. Concurrently, growth rates and yields in laterinitiated tillers were reduced in relation to their leaf areas.Stem growth rate was proportionately increased more than paniclegrowth rate in the longer DVP treatments and this, combinedwith a longer duration of stem growth, resulted in greater stemdry matter at maturity and, therefore, in reduced harvest index.     

An Association between Flowering and Reduced Stomatal Sensitivity to Water Stress in Pearl Millet [Pennisetum americanum (L.) Leeke]

Stomatal sensitivity to water stress was investigated in pearlmillet [Pennisetum americanum (L.) Leeke] in relation to stageof plant development, leaf water status and ABA content by samplingplants at midday. For the same leaf water potential (), droughtedplants with emerged panicles were found to have a greater leafconductance (g_L), indicative of greater stomatal opening, thanplants sampled prior to panicle emergence. The difference betweensuch flowering (F) and non-flowering (NF) plants in at stomatalclosure was estimated to be at least 0.6 MPa. This differencewas considered unlikely to be the result of differential bulkleaf osmotic adjustment, and for most samples from both F andNF plants, bulk leaf turgor potential (_p) was estimated to bezero. Stomatal closure in NF plants was associated in two genotypes(BJ 104 and line 112) with higher leaf ABA levels. Differencesin ABA levels between F and NF plants were, however, smalleror absent in genotypes Serere 39 and B282. These genotypes wereat lower than BJ 104 and line 112 when sampled and showed smallerdifferences between F and NF plants in conductance. Lower ABA levels in F plants are ascribed either to effectsof leaf ageing or to effects of flowering on ABA content ofthe leaf. Significant differences in g_L in the absence of differencesin ABA content are taken to imply changes in stomatal sensitivityto the hormone or in its access to the stomatal complex. Pennisetum americanum (L.) Leeke, pearl millet, flowering, stomata, water stress, abscisic acid     

Stomatal responses of pearl millet (Pennisetum americanum [L.] Leeke) to leaf water status and environmental factors in the field

Abstract. Factors affecting stomatal conductance (g₁) of pearl millet ( Pennisetum americanum [L.] Leeke), cultivar BJ 104, were examined in the field in India during the dry season.
Diurnal changes in g₁ were evaluated for upper expanded leaves at flowering on two occasions using plants subjected to varying degrees of water stress. Except for the most severely stressed treatment, diurnal changes in g₁ closely matched changes in irradiance ( I ), the promotive effect of which largely overcame opposing influences on g₁ of increasing atmospheric vapour pressure deficit, and decreasing leaf water and turgor potentials (Ψ, Ψ_p).
Two main effects of water stress on g₁ were evident: (i) a decrease in the amplitude of the mid-day peak in g₁, and (ii) a decrease in the time over which high g₁ was maintained, resulting in early (mid-day) closure and hysteresis in the relationship between g₁ and I .
Leaf conductance was greatest for upper leaves and decreased down the canopy. At equivalent depths in the canopy g₁ was higher in flowering than in photoperiodically-retarded plants of the same age. The magnitude of water stress-induced stomatal closure increased down the plant, and was more marked in retarded than in flowering plants.
Within individual stress treatments Ψ of upper leaves decreased linearly as transpiration flux increased. It is concluded that stomatal behaviour of upper leaves of pearl millet at flowering largely operates to maximize assimilation rather than to minimize water loss.     

Stomatal Responses of Pearl Millet (Pennisetum americanum (L.) Leeke) Genotypes, in Relation to Abscisic Acid and Water Stress

Stomatal responses to water stress and to applied (±)-abscisicacid (ABA) were examined in genotypes of pearl millet (Pennisetumamericanum (L.) Leeke) known to differ in amounts of endogenousABA accumulating during drought. In both a pot and a field experiment,Serere 39, a genotype with a high capacity to accumulate ABA,showed a higher stomatal sensitivity to water stress than didthe ‘low’ ABA accumulator, BJ 104. In the fieldexperiment, a third genotype, B282, accumulating least amountsof ABA, also had the lowest stomatal sensitivity to water stress. There were no significant differences between these genotypesin stomatal response to applied (±)-ABA, or in the relationshipsbetween leaf conductance and levels of endogenous ABA. It isconcluded that the differences in accumulation of endogenousABA by these genotypes of pearl millet are of functional significance,and that endogenous ABA generated during a water stress whichdevelops over days or weeks mediates stomatal responses to suchstress.     

The Effect of Flowering on Stomatal Response to Water Stress in Pearl Millet (Pennisetum americanum [L.] Leeke)

Stomata on upper leaves of drought-stressed pearl millet (Pennisetumamericanum [L.] Leeke) crops were more open in flowering (F)than in pre-flowering (PF) plants. This was not due to differencesin leaf water potential (). Stomata of PF plants closed when fell to about –1.7 MPa, while on F plants stomata closedonly when approached –2.3 MPa. Osmotic adjustment did not account for these differences asrelations between turgor potential (_P) and were similar inF and PF plants. While stomata of PF plants closed as W becamezero, in F plants stomata remained open even after bulk leafturgor was lost. Differences between F and PF plants were not explained by differencesin age of leaves sampled. However, leaves of water-stressedPF plants had higher levels of abscisic acid (ABA) than leavesof F plants, despite similarities in water status. From theseresults and from relationships between g_L and stage of panicledevelopment, it is concluded that the tendency of stomata toremain open despite water stress and loss of bulk leaf _P isrelated to the presence of an emerged panicle. Hypotheses whichaccount for this effect are discussed. Key words: Pennisetum americanum [L.] Leeke, Pearl millet, Flowering, Stomata, Water stress, Abscisic acid     

Genotypic Variation in Pearl Millet (Pennisetum americanum (L.) Leeke), in the Ability to Accumulate Abscisic Acid in Response to Water Stress

Variation between genotypes in the ability to accumulate abscisicacid (ABA) in response to water stress was investigated in pearlmillet (Pennisetum americanum (L.) Leeke). Using a detachedleaf test a more than four-fold variation in accumulation capacitywas observed amongst a set of 16 genotypes grown in a controlledenvironment. Two genotypes which contrasted in their accumulationcapacity, BJ 104 and Serere 39 (the latter accumulating mostABA), maintained the difference over a range of leaf water contentsand potentials. Some of the genotypes were grown in the field in the semi-aridtropics with and without irrigation, and sampled for ABA content.In two experiments, substantial genotypic variation in ABA accumulationwas observed, which could not be attributed to differences inleaf water potential (). In a field experiment comparing threegenotypes (Serere 39, BJ 104, and B282), differences in ABAaccumulation were also shown to be largely independent of genotypicdifferences in turgor potential (_p). For a set of six of the genotypes, the amounts of ABA accumulatingin leaves of intact, droughted plants, in the field, when adjustedfor differences in , were found to be significantly (P<0.05)correlated with amounts of ABA accumulated in detached, water-stressedleaves. It is concluded that the detached leaf test adequatelyreflects the ability of pearl millet genotypes to accumulateABA under field conditions.     

Stomatal Response to Water Stress and its Relationship to Bulk Leaf Water Status and Osmotic Adjustment in Pearl Millet (Pennisetum americanum [L.] Leeke)

The water potential () at which stomata completed closure (_8Lmin)was determined for pearl millet (Pennisetum americanum [L.]Leeke) at two growth stages by monitoring changes in leaf conductance(g_L) and following shoot detachment. Leaf water status wasevaluated concurrently using a pressure-volume (P-V) technique. In a pot experiment with young vegetative plants, _8Lmin closelyapproximated to the estimated at zero turgor (_u) both for controland for drought-conditioned plants which had osmotically adjusted.However, for penultimate leaves of field-grown flowering plants,_8Lmin was found to be 0.61 (irrigated plants) and 0.87 (droughtedplants) MPa below _u. In drought-stressed field-grown plants,osmotic adjustment (characterized by a decrease in solute (osmotic)potential (_s ) at both full hydration and zero turgor) was insufficientto maintain a positive bulk leaf turgor potential (_p) once had declined to below about -1.5 MPa. It is suggested that localizedadjustment by the stomatal complex in response to environmentaldifferences, leaf ageing and/or ontogenetic change, is responsiblefor the uncoupling of stomatal from bulk leaf water status. Key words: Stomata, Water stress, Pennisetum americanum     

Diurnal Changes in Endogenous Abscisic Acid in Leaves of Pearl Millet (Pennisetum americanum (L. ) Leeke) under Field Conditions

Diurnal variation in leaf abscisic acid (ABA) content was investigatedin pearl millet (Pennisetum americanum (L. ) Leeke) growingin the field in the semi-arid tropics and subjected to varyingdegrees of water stress. There was a two- to three-fold change in ABA content duringthe photoperiod in three groups of ‘severely’ stressedplants of the genotype BJ 104. Maximum ABA occurred mid-morning(1030 h). ABA levels then declined to a minimum at 1500 h. Changesin ABA content of ‘moderately’ stressed and fullyirrigated plants were smaller, but still significant. Though,when averaged over the day, levels of ABA of the five groupswere positively related to the degree of water stress, relationshipsbetween ABA concentration and total water () or turgor (_p) potentialsvaried considerably with time of sampling. Within groups, changesin ABA contents during the day were not always accounted forby changes in or _p. Temporal changes in leaf ABA content similar to those foundin BJ 104, and largely unrelated to , were observed in the genotypesSerere 39 and B282 in a subsequent year. Leaf ABA content of droughted plants (BJ 104) did not declineappreciably overnight despite a marked increase in . However,a large reduction in ABA content with increase in did occurfollowing heavy rainfall. Diurnal changes in stomatal conductance (g₁) of BJ 104 couldnot be simply accounted for by temporal changes in total leafABA content, even when allowance was made for effects of irradianceand other environmental variables on g₁. It is suggested thatthe sensitivity of stomata to ABA, or accessibility of the hormoneto the stomatal complex, changes during the day.     

Leaf Abscisic Acid Content and Recovery from Water Stress in Pearl Millet (Pennisetum americanum [L.] Leeke)1

Water potential () and abscisic acid (ABA) content were measuredin leaves of drought-stressed, field-grown plants of pearl millet(Pennisetum americanum [L.] Leeke) during rehydration, initiatedeither in response to a diurnal night-time reduction in evaporativedemand, or to irrigation. Overnight rehydration, manifestedas a substantial increase in , was not accompanied by any reductionin ABA levels. In contrast, an increase in following irrigationresulted in an appreciable reduction in ABA content. Such reductionwas, initially, only partial when field plants were rewateredat dusk, but was rapid and complete within 3 h when irrigationwas applied mid-afternoon. The possibility that light or temperature changes might haveprevented loss of ABA during night-time rehydration was investigatedin pot experiments. At similar air temperatures, young pot-grownplants rehydrated more rapidly, and ABA levels fell more quickly,in darkness than in light. The onset of rehydration and lossof ABA in darkness were delayed by low (20 ?C) compared withhigh (28 ?C) temperature, though after an initial lag, ratesof both processes at 20 ?C were similar to rates at 28 ?C. Neitherlight nor temperature affected the relationship between ABAcontent and *. Key words: Abscisic acid, Rehydration, Pennisetum americanum     

Osmotic adjustment to water stress in pearl millet (Pennisetum americanum [L.] Leeke) under field conditions

Abstract. Osmotic adjustment, a mechanism whereby plants maintain positive turgor despite low water potential (ψ), was investigated in pearl millet ( Pennisetum americanum [L.] Leeke) in three types of field experiment at Hyderabad, India:

• (1)

  Osmotic adjustment during the growing season was evaluated by comparing solute potential (ψ_s) of leaves taken at midday from irrigated and droughted plots and allowed to rehydrate in the laboratory. The degree of seasonal adjustment was also estimated by comparing observed values of ψ_s in the field with those expected if ψ_s decreased solely in proportion to water loss. Both types of assessment indicated the maximum seasonal adjustment to be about 0.2 MPa. The cultivars BJ 104 and Serere 39 differed in their capacity to adjust osmotically over the season; Serere 39 was least able to osmoregulate.
• (2)

  Measurements of diurnal variations in ψ and ψ_s in BJ 104 revealed osmotic adjustment during the afternoon hours. At a given value of ψ, turgor (ψ_p) was about 0.1 MPa higher in irrigated, and over 0.2 MPa higher in droughted plants, in the afternoon, than in the morning.
• (3)

  Osmotic adjustment of different leaves within the canopy was investigated. Upper leaves had lower ψ than basal leaves. Differences in ψ were matched by gradients in ψ_s, so that turgor was similar for all leaf layers.