Effect of Matric-Priming Duration and Priming Water Potential on Germination of Four Grasses

In this study, a matric-potential control system was used todetermine the effect of matric-priming duration and primingwater potential on the germination response of Bouteloua curtipendula(Michx.) Torr., Cenchrus ciliaris L., Eragrostis lehmannianaNees, and Panicum coloratum L. Seeds were primed at water potentialsof –1·5 to –7·7 MPa for up to 14 d.Optimum germination generally occurred in treatments primedat high water potential for the shortest period. Germinationof seeds primed at lower water potential and for longer periodsexhibited a negative germination response relative to the control.Seeds were not redried after the priming treatment. Seed-wateruptake measurements suggest that a reduction in the lag timeof imbibition accounted for at least some germination-rate enhancementin the positive-priming treatments Key words: Germination, matric-priming, imbibition     

Effects of Temperature and Water Potential on Germination, Radicle Elongation and Emergence of Mungbean

Controlled environment experiments were performed to determinethe effects of temperature and water potential on germination,radicle elongation and emergence of mungbean (Vigna radiata(L.) Wilczek cv. IPB-M79-17-79). The effects of a range of constant temperatures (15–45°C) and water potentials (0 to –2.2 MPa) on germinationand radicle elongation rates were studied using an osmoticumtechnique, in which seeds were held against a semi-permeablemembrane sac containing a polyethylene glycol solution. Linearrelationships were established between median germination time(Gt₅₀) and water potential at different temperatures, and betweenreciprocal Gt₅₀ (germination rate) and temperature at differentwater potentials. Germination occurred at potentials as lowas –2.2 MPa at favourable temperatures (30–40 °C),but was fastest at 40 °C when water was not limiting, withan estimated base temperature (T_b) of about 10 °C. Subsequentradicle elongation, however, was restricted to a slightly narrowertemperature range and was fastest at 35 °C. The conceptof thermal time was used to develop an equation to model thecombined effects of water potential and temperature on germination.Predictions made using this model were compared with the actualgermination obtained in a related series of experiments in columnsof soil. Some differences observed suggested the additionalimportance of the seed/soil/water contact zone in influencingseed germination in soil. Seedling emergence appeared to reflectfurther the radicle elongation results by occurring within anarrower range of temperatures and water potentials than germination.Emergence had an estimated T_b of 12.6 °C and was fastestat 35 °C. A soil matric potential of not less than about–0.5 MPa at sowing was required to obtain 50% or moreseedling emergence. Key words: Germination, temperature, water potential     

The Effect of Soil Water and Aeration on Seed Germination

The time rate of germination and the final germination percentageof Oryzopsis holciformis decreased with increasing water stress.The optimum matric potential for germination was–0.005bar in coarse sand and –0.5 bar in sandy loam soil. Thisdiscrepancy was explained by changes in the rate of water-supplyto the seed, as determined by the area of contact between seedand germination medium, and by the hydraulic conductivity ofthe medium. At high soil moisture potentials germination also decreased.Such a decrease was not found at equivalent osmotic potentials.It seems that this decrease in germination was brought aboutby the thickening of the water films around the seeds, whichinterfered with oxygen diffusion. This assumption was supportedby determinations with Pt electrodes, and by previous work ongermination at lowered oxygen concentrations.     

Nitrate Stimulation of Mobilization of Seed Reserves in Temperate Cereals: Importance of Water Uptake

Relationships between nitrate (NO^-₃) supply, uptake and assimilation,water uptake and the rate of mobilization of seed reserves wereexamined for the five main temperate cereals prior to emergencefrom the substrate. For all species, 21 d after sowing (DAS),residual seed dry weight (d.wt) decreased while shoot plus rootd.wt increased (15–30%) with increased applied NO^-₃concentrationfrom 0 to 5–20 mM . Nitrogen (N) uptake and assimilationwere as great with addition of 5 mM ammonium (NH⁺₄) or 5 mMNO^-₃but NH⁺₄did not affect the rate of mobilization of seedreserves. Chloride (Cl^-) was similar to NO^-₃in its effect onmobilization of seed reserves of barley (Hordeum vulgare L.).Increased rate of mobilization of seed reserves with additionalNO^-₃or Cl^-was associated with increases in shoot, root and residualseed anion content, total seedling water and residual seed watercontent (% water) 21 DAS. Addition of NH⁺₄did not affect totalseedling water or residual seed water content. For barley suppliedwith different concentrations of NO^-₃or mannitol, the rate ofmobilization of seed reserves was positively correlated (r >0.95)with total seedling water and residual seed water content. Therate of mobilization of seed reserves of barley was greaterfor high N content seed than for low N content seed. Seed watercontent was greater for high N seed than for low N seed, 2 DAS.Additional NO^-₃did not affect total seedling water or residualseed water content until 10–14 DAS. The effects of seedN and NO^-₃on mobilization of seed reserves were detected 10and 14 DAS, respectively. It is proposed that the increasedrate of mobilization of seed reserves of temperate cereals withadditional NO^-₃is due to increased water uptake by the seedlingwhile the seed N effect is due to increased water uptake bythe seed directly. Avena sativa L.; oat; Hordeum vulgare L.; barley; Secale cereale L.; rye; xTriticosecale Wittm.; triticale; Triticum aestivum L.; wheat; nitrate; seed; germination; seed reserve mobilization     

Influence of Soil Water Matric Potential and Hydraulic Conductivity on the Germination of Rape (Brassica napus L.)

The influence of matric potential and hydraulic conductivityon the water absorption and germination of rape (Brassica napusL.) was investigated by comparative studies in three seed-soilwater systems. In all systems the rate of germination and totalgermination were substantially retarded with decreasing matricpotential. Hydraulic conductivity of the soil system was demonstratedto be a limiting factor in the germination process. The effectsof matric potential and hydraulic conductivity on water absorptionfollowed closely those demonstrated for germination. The influenceof matric potential was shown to be comparable to that of osmoticpotential provided consideration was given to the hydraulicproperties of the soil-seed system.     

Water Uptake and Protein Synthesis in Germinating Wheat Embryos under the Osmotic Stress of Polyethylene Glycol

The effect of osmotic stress on wheat-seed germination was testedby imbibition in aqueous polyethylene glycol solutions at differentconcentrations. The experiments were designed to allow blockingand the subsequent recovery of germination by 12 h or 24 h pre-imbibitionof seeds in osmoticum, followed by transfer to water. Seedswere alternatively presoaked in water for 12 or 24 h, then transferredto polyethylene-glycol solutions to study the induced blockingof germination. Water content and [³H]leucine incorporationinto embryo tissues (as a measure of in vivo protein synthesis)were determined over a 48-h imbibition period. A close relationshipwas established overall between hydration status and proteinsynthesis rate. Osmotic stress seems to have a strong influenceupon the quantitative synthesis of proteins, suggesting thatthis biochemical activity is associated with the regulationof the germination process. Triticum durum, embryo, osmotic stress, water uptake, protein synthesis     

Water uptake and distribution in non-dormant and dormant wild oat (Avena fatua L.) caryopses

The influence of seed coat modification and light quality onwater uptake and distribution in caryopses of dormant and non-dormantlines of wild oat (Avena fatua L.) was determined using NMRmicroimaging. Non-dormant seeds absorbed water more rapidlythan dormant seeds during imbibition on distilled water. Thiseffect was detected first in the embryo-scutellar region (8h) and later in the proximal endosperm (12 h). Cutting the testaand pericarp close to the embryo or scarification with KOH promotedrapid embryo/scutellum hydration and germination. Cutting atthe middle part of the caryopsis did not enhance embryo hydrationnor did it greatly improve germination. The sensitivity of waterdistribution to the phytochrome germination effect was examined.Significant differences in imbibitional water uptake by embryos-scutellumtissue were detected by 18 h following red-light (germinationpromoter) compared with far-red (germination inhibitor) treatment.The results indicated that both the rate and the sequence ofembryo/scutellum hydration were important in initiating germinationin dormant seeds. A refinement of the model that describes waterimbibition in wild oat seeds during the early stages of germinationis discussed. Key words: Water uptake, water distribution, Avena fatua, seed coat modification, light quality, dormant and non-dormant seeds     

The Relation of Water Absorption by Wheat Seeds to Water Potential

The progress of water absorption by wheat grains was studiedby supplying water in the vapour phase, at controlled potentials. At a potential of –250 metres of water, the curve forwater uptake against time shows exponential approach to equilibriummoisture content. Living and dead seeds behave similarly untilgermination effects are apparent. Water uptake in the earlystages is probably due to physical rather than physiologicalprocesses. When germination occurs, it causes an exponentialincrease in the rate of water uptake. At higher potentials, up to zero, the uptake curves for deadseeds depart from the simple exponential relationship; in additionto the exponential component, there is a component the rateof which increases with time to a constant rate. The first componentmay represent the physical process of imbibition by the starch,and the second the initiation and progress of starch hydrolysis. A parameter a of the formula derived for the curves is interpretedas representing the diffusivity of water vapour in the seedmaterial, depending upon the physical properties and dimensionsof the seed.     

Short-term Effects of Ferulic Acid on Ion Uptake and Water Relations in Cucumber Seedlings

Ferulic acid (FA) is commonly found in soils and is consideredan allelochemical. Studies have suggested that FA and otherphenolic acids decrease plant growth in part by decreasing theabsorption of mineral nutrients and water. However, no studieshave examined these parameters in a single experimental systemto investigate how FA affected both ion uptake and plant-waterrelations in whole plants. Using intact cucumber (Cucumis sativusL. cv. Early Green Cluster) seedlings, we examined short-termeffects of FA on ion uptake kinetics, transport promoters andinhibitors, and water relations as indicated by a pressure-volumeanalysis. We found that after 3 h of treatment, 200 µMFA inhibited net ion uptake, particularly NO₃, and promotednet K⁺ efflux from seedling roots. The addition of fusicoccin,a K⁺ transport promoter, counteracted the inhibitory effectof FA on net K⁺ uptake. Concurrent treatment of seedlings withFA and tetraethylammonium, a channel-blocking salt, reducedaverage K⁺ efflux by 66%. Treatment of seedlings with FA alsodecreased leaf water potential (₁ and turgor pressure (P_T).However, decreased ₁ and P_T were not caused by changes in theosmotic properties of the symplast or stomatal conductance.A decrease in water absorption is a likely explanation for theloss of P_T observed. The results of our experiments indicatethat both ion uptake and plant-water relations can be adverselyaffected by FA. Key words: Cucumis sativus, ferulic acid, allelopathy, ion uptake, water relations     

Water Relations of Seed Development and Germination in Muskmelon (Cucumis melo L.): VII. INFLUENCE OF AFTER-RIPENING AND AGEING ON GERMINATION RESPONSES TO TEMPERATURE AND WATER POTENTIAL

The effects of storage conditions on the germination of developingmuskmelon (Cucumis melo L.) seeds were tested to determine whetherafter-ripening is required to obtain maximum seed vigour. Seedswere harvested at 5 d intervals from 35 (immature) to 60 (fullymature) days after anthesis (DAA), washed, dried, and storedat water contents of 3·3 to 19% (dry weight basis) at6, 20, or 30°C for up to one year. Germination was testedin water and in polyethylene glycol 8000 solutions ( –0·2to –1·2 MPa osmotic potential) at 15, 20, 25 or30°C. Germination percentages and rates (inverse of meantimes to radicle emergence) were compared to those of newlyharvested, washed and dried seeds. For 40 and 60 DAA seeds,one year of storage at 20°C and water contents <6·5%significantly increased germination percentages and rates at20°C, but had little effect on germination at 25 and 30°C.Storage reduced the estimated base temperature (T_b) and meanbase water potential (_b) for germination of both 40 and 60 DAAseeds by approximately 5°C and 0·3 MPa, respectively.Immature 35 DAA seeds showed the greatest benefit from storageat 3 to 5% water content and 30°C, as germination percentagesand rates increased at all water potentials (). Storage underthese same conditions had little effect on the germination ofmature seeds in water, but increased germination percentagesand rates at reduced 's. Accelerated ageing for one month at30°C and water contents from 15 to 19° increased germinationrates and percentages of mature seeds at reduced 's, but longerdurations resulted in sharp declines in both parameters. Immatureseeds lost viability within one month under accelerated ageingconditions. An after-ripening period is required at all stagesof muskmelon seed development to expand the temperature andwater potential ranges allowing germination and to achieve maximumgerminability and vigour. Post-harvest dormancy is deepest atthe point of maximum seed dry weight accumulation and declinesthereafter, both in situ within the ripening fruit and duringdry storage. Key words: Muskmelon, Cucumis melo L., seed, development, dormancy, germination, vigour, after-ripening     

Seed Water Relations and the Regulation of the Duration of Seed Growth in Soybean

Soybean [Glycine max (L.) Merrill] seeds and cotyledons weregrown in an in vitro culture system to investigate the relationshipsbetween cell expansion (net water uptake by the seed) and drymatter accumulation. Seeds or cotyledons grown in a completenutrient medium containing 200 mol m^–3 sucrose continueddry matter accumulation for up to 16 d after in planta seedsreached physiological maturity (maximum seed dry weight). Seedor cotyledon water content increased throughout the cultureperiod and the water concentration remained above 600 g kg^–1fresh weight. These data indicate that the cessation of seeddry matter accumulation is controlled by the physiological environmentof the seed and is not a pre-determined seed characteristic.Adding 600 mol m^–3 mannitol to the medium caused a decreasein seed water content and concentration. Seeds in this mediumstopped accumulating dry matter at a water concentration ofapproximately 550 g kg^–1. The data suggest that dry matteraccumulation by soybean seeds can continue only as long as thereis a net uptake of water to drive cell expansion. In the absenceof a net water uptake, continued dry matter accumulation causesdesiccation which triggers maturation. Key words: Glycine max (L.) Merrill, solution culture, duration of seed growth, water content, dry matter accumulation     

Water Transport during Acid-Induced Growth of Excised Hypocotyl Sections of Vigna unguiculata under Xylem Perfusion

Elongation growth of abraded hypocotyl sections of Vigna unguiculataunder xylem perfusion was markedly promoted a few minutes afterthe application of an acid aerosol generated from a solutionof HCl. At the beginning of the acid-induced growth, intracellularpressure (Pⁱ) began to decrease and the membrane potential betweenthe symplast and the xylem apoplast (V^px) began to depolarize.Subsequently, Pⁱ and V^px remained at a reduced level and a depolarizedlevel, respectively, while the promotion of elongation growthcontinued for more than 4 hours. The electrogenic componentof the xylem membrane potential (V^px_act) gradually increasedto about twice that before acid treatment. There was a closecorrelation between the enhanced growth and the decrease inintracellular pressure within 30 min after application of acidbut little correltion after 60 min. By contrast, there was littlecorrelation between the promotion of growth and the activityof the xylem pump after 30 min while a close correlation wasobserved after 60 min. It is inferred that the acid-induced activation of water uptakeconsists of two major processes, in series, that are drivenby different forces: the rapid uptake of water for more than30 min, driven by hydrostatic force generated by loosening ofcell walls; and a long-lasting enhancement of water uptake forat least 4 h, which is driven by osmotic force that is generatedby the canal system within the xylem. (Received October 17, 1994; Accepted January 23, 1995)     

Germination and Early Growth of Plants Studied Using Neutron Radiography

Seed swelling, germination, root extension, lateral root initiationand shoot growth were studied in soils of different water contents,using non-destructive, serial neutron radiography. Seeds fromthree varieties of soya beans (Glycine max L.) and one varietyeach of maize (Zea mays L.) and vetch (Vicia sativa L.) wereused. The seeds germinated when they had increased in size bya certain amount, if germination is taken as the time when theradicle first appears. The rate at which roots and shoots extendalso depend on soil water content. Glycine max L., Vicia sativa L., Zea mays L., Soya bean vetch, maize, seed germination, root extension, lateral root initiation, neutron radiography     

Effect of Water Potential and Temperature on the Germination of Four Species of African Savanna Trees

The germination responses of seeds from the African tree speciesColophospermum mopane, Combretum apiculatum, Acacia tortilisand Acacia karroo under varying regimes of temperature and waterstress (induced by incubation in PEG 8000) are reported Withthe exception of Combretum (at –0.14 and –0.29 MPa)and Colophospermum (at –0.29 MPa), incubation in PEG decreasedthe maximum achieved germination percentage (90–100% forall species), but did not extend the germination lag (exceptin Combretum) or affect the time required to reach maximum germinationCombretum and Colophospermum were found to germinate under thewidest range of temperatures and water potentials, for example,as strongly negative as –1.0 MPa at 20 and 30 °C,respectively These seeds also showed greater or equivalent hypocotylelongation in PEG solutions creating potentials of –0.14,–0.29 or –0.51 MPa when compared with seeds germinatedin water, indicating an additional stress adaptation Acaciaspecies showed progressive reduction in germination rates andradicle elongation in response to decreasing water potentialExperiments giving pre-imbibition treatments in water priorto transfer to PEG solutions showed that both Acacia speciesgerminated at approximately 90% if given such pre-treatmentand less than 10% if transferred directly to PEG It is concludedthat the most stress-adapted species studied are Colophospermummopane and Combretum apiculatum, a finding generally correlatedwith the growth habit of these trees Colophospermum mopane, Combretum apiculatum, Acacia tortilis, Acacia karroo, germination, water stress, Zimbabwe     

Heterogeneity in Water Availability Alters Cellular Development and Hydraulic Conductivity along Roots of a Desert Succulent

Plants of the desert succulent Agave deserti were grown in partitionedcontainers to determine whether heterogeneity in soil moistureleads to differences in cellular development and hydraulic conductivityalong individual roots. Roots from containers with a dry distalcompartment (furthest from the shoot), a wet middle compartment,and a dry proximal compartment had distal regions (includingthe root tips) that were more suberized and lignified in theendodermis and adjacent cell layers than were root regions fromthe wet middle compartment. Proximal root regions about 40 mmfrom the succulent shoot base were also relatively unsuberized,suggesting that both external and internal supplies of waterdelayed tissue maturation. Root segments from wet middle compartmentsand from dry proximal compartments had higher hydraulic conductivitythan did the more suberized root segments from dry distal compartments.Unlike distal root segments from wet compartments, segmentsfrom dry compartments suffered no decrease in hydraulic conductivityafter immersion in mercuric chloride, suggesting that aquaporinactivity diminished for roots during drought. The possible closureof water channels could help limit root water loss to a dryingsoil. The delayed development of suberized cell layers may allowroot regions to maximize water uptake from wet soil patches(such as under rocks), and the relatively immature, absorptiveroot region near the base of the shoot may help A. deserti capturewater from a briefly wetted surface soil. Copyright 2000 Annalsof Botany Company Agave deserti, root plasticity, water uptake, aquaporins, suberization, endodermis, divided pots.     

The Response of Oxygen and Carbon Dioxide Exchanges and Root Activity to Short Term Water Stress in Soybean

The gross and net O₂ evolution together with O₂ uptake, CO₂assimilation, transpiration, shoot dark respiration, root respirationand ion uptake of a soybean plant were studied during 19 d whichincluded two periods of water stress. O₂ uptake was measuredusing ¹⁸O₂ as a tracer. Short term water stress induced immediateand lasting effects: (1) reduction of light interception bywilting, (2) limitation of the total reducing equivalent producedby the electron transport chain, (3) decrease of stomatal conductancereducing both losses of water and the entry of CO₂ for assimilation,(4) relative stimulation of O₂ uptake. The ratio of O₂ uptaketo CO₂ assimilation changed from 1.0 before stress to 1.4 forseveral days after. Root respiration was less affected by thestress than ion uptake and shoot gas exchanges. Key words: Photosynthesis, Photorespiration, Transpiration, Shoot and root respiration, Ion uptake, Water stress, Glycine max. L.     

Water Import Rate in Tomato Fruit: A Resistance Model

A model of the water import rate in tomato fruit is proposed.It compares the fruit to a hollow sphere (P) with external radiusR and internal radius R_G, corresponding to pericarp, and containingan internal spherical part (G). The pathway limiting water inputrate at any point I at a distance r from the fruit centre wasassumed to be proportional: (a) in P, to the length of the arcwhich has a radius r and which goes from I to the pedicel extension;(b) in G, to r. The water input rate at I was modelled basedon a law similar to Darcy's law which takes into account thedifference between the water potential at entry of fruit andthe water potential at point I. This latter potential was thesum of fruit osmotic potential and pressure potential due toresistance of tissue to deformation. This potential was proportionalto R-r or R_G -r. The model was expressed at fruit level by alaw such that water mass imported per unit time per unit surfacearea of fruit (f_rw) was a linear function of R. The model wascompared to linear regressions of this rate in terms of R whichhad been found during fruit swelling from published results,and which were obtained at different values of nutrient solutionsalinity. The results suggested that water input in tomato fruitis conditioned by passive forces depending on fruit size.Copyright1994, 1999 Academic Press Fruit, growth, model, resistance, salinity, size, tomato, transfer, water     

The Use of Coleoptile Responses to Water Stress to Differentiate Wheat Genotypes for Osmoregulation, Growth and Yield

Differences in osmoregulation between genotypes of wheat (Triticumaestivum L.) were identified by measuring responses of coleoptilesto water stress. Seeds were germinated in Petri dishes on germinationpads. Water stress was applied when coleoptiles were 1 cm longby either adding polyethylene glycol 6000 (20% w/w) or by removingthe lids of the dishes and allowing the pads to dry by evaporationfor a period of 2 d, or from the beginning of germination byvarying the amount of water in the germination pads. Measurementsof the water and osmotic potentials of shoots and roots (usingthermocouple psychrometers) were used to estimate turgor pressures.Wheat genotypes showed the same segregation for shoot and rootlengths and turgor pressures as they had done in previous studiesfor flag-leaf osmoregulation and yield. It was concluded thatobservations of the responses of coleoptile length to waterstress could be used in plant-breeding applications to identifydifferences in osmoregulation between genotypes of wheat. Triticum aestivum, osmoregulation, turgor maintenance, coleoptile growth, root, selection criterion     

Altered Synthesis and Composition of Cell Wall of Grape (Vitis vinifera L.) Leaves during Expansion and Growth-Inhibiting Water Deficits

The rate and composition of cell wall polysaccharide synthesisduring development and growth-inhibiting water deficits wereinvestigated in leaves of grape (Vitis vinifera L.). The rateof leaf expansion was monitored as plant water status was manipulatedby modulating the supply of irrigation water to potted plantsover several days. The corresponding wall synthesis was determinedby incubating leaf tissue with [¹⁴C]glucose and quantifyingincorporation into wall components. Samples were obtained fromrapidly expanding and mature leaves before, during, and following(recovery from) moderate water deficits. Uptake was approximately2-fold greater for mature leaf tissue than for rapidly expandingtissue at both high and low water status. In contrast, incorporationinto cell wall polysaccharides was 18 to 41% (under low andhigh water status) of uptake in expanding leaves but less than4% in mature tissue. Incorporation of precursor into wall polysaccharideswas insensitive to plant water status in mature leaves, butwas inhibited to less than 50% of well-watered controls in expandingleaves at low water potential. Incorporation of label into cellulose,uronic acid, and neutral sugar fractions was differentiallyaffected by water deficits, with cellulose synthesis apparentlyexhibiting the greatest sensitivity to low water status. Afterrewatering, growth, as well as uptake and incorporation of labelrecovered, although the latter did not attain prestress rates.The results indicate a high sensitivity of wall polysaccharide(particularly cellulose) synthesis to growth-inhibiting waterdeficits. ¹ Supported by United States Department of Agriculture, CompetitiveResearch grant GAM 8502539. (Received November 15, 1989; Accepted January 17, 1990)     

Photoinhibition of radish (Raphanus sativus L) seed germination: control of growth potential by cell-wall yielding in the embryo

The biophysical mechanism underlying photoinhibition of radish (Raphanus sativus L.) seed germination was investigated using three cultivars differing in sensitivity to continuous irradiation with far-red light (high-irradiance reaction of phytochrome). Sensitivity of germination to the inhibitory action of light was assessed by probing germination under osmotic stress (incubation in media of low water potentials adjusted with polyethylene glycol 6000) and expressed in terms of ‘germination potential’ (positive value of the water potential at which germination is inhibited by 50%). Far-red light decreases the germination potential to various degrees in the different cultivars, reflecting the light-sensitivity of germination in water. Removal of the seed coat increases the germination potential by a constant amount in darkness and light. It is concluded that germination depends on the expansive force of the embryo which can be drastically diminished by far-red light. Seed-coat constraint and expansive force of the embryo interact additively on the level of the germination potential. Photoinhibition of germination was accompanied by an inhibition of water uptake into the seed. Analysis of seed water relations showed that osmotic pressure and turgor assumed higher levels in photoinhibited seeds, compared to seeds germinating in darkness, while the water potential was close to zero under both conditions. Far-red light produced a shift (to less negative values) in the curve relating water-uptake rate to external water potential, i.e. a reduction in the driving force for water uptake. It is concluded that photoinhibition of germination results from the maintenance of a high threshold of cell-wall extensibility in the embryo.