The Effect of Water Stress on Floral Characters, Pollination and Seed Set in White Clover (Trifolium repens L.)

White clover plants were subjected to either short-term developingwater stress or long-term stable levels of water deficit. Thehort-term stress reduced plant water status to about –2·0MPa over 16 d. The long-term stress was less severe, but wassustained for several weeks. Long-term water stress promotedthe production of inflorescences. However, water stress alsoincreased floret abortion and the premature death of whole flowerheads. The number of ovules per floret was decreased by waterstress. The most striking effect of both long- and short-term waterdeficit was to reduce pollen viability measured with the fluorochromaticassay. This was not an artefact of assay conditions. The pollenfrom water-stressed flower heads was not reversibly dehydrated;it did not score at similar viability to controls after incubationin conditions which hydrate pollen. In addition, the pollenfrom water-stressed plants lost viability more rapidly thanpollen from well-watered plants after removal from the flowerhead. The consequences of reduced pollen viability on seed set wereinvestigated by hand-crossing within and between groups of plantsmaintained for several weeks at three levels of water supply.Flower heads pollinated with pollen from water-stressed plantsset fewer seeds per floret than those pollinated with controlpollen. Key words: Trifolium repens, white clover, water stress, floral characters, seed set     

The Extent and Pattern of Osmotic Adjustment in White Clover (Trifolium repens L.) During the Development of Water Stress

White clover plants were subjected to water stress followingthe cessation of watering. As a water deficit developed, waterand osmotic potentials were measured in stolon tips, leavesfrom the stolon tip and leaves from the plant crown. Pressurepotentials were calculated. Pressure potential was maintainedin stolon tips even when water potential fell to around –2·0MPa. In contrast, pressure potential in leaves fell rapidlyas water stress developed. Total amino acid and potassium levels were largely unaffectedin both stolon tips and leaves. Water-soluble carbohydratesand proline accumulated during water stress. The increase inproline level in leaves did not follow the same pattern as thatin stolon tips, although toward the end of the water stressperiod the level had increased by a similar extent in both partsof the plant. Additionally, pressure potential and osmotic potentialappeared to be significantly related to proline content in stolontips. No such relationship was found for leaves. The role ofproline in osmotic adjustment is discussed. Trifolium repens L. cv. Olwen, white clover, water stress, osmotic adjustment, proline     

Influence of Water Stress on Flowering and Seed Production of Macroptilium atropurpureum cv. Siratro

Macroptilium atropurpureum cv. Siratro was grown in large soilbeds with a constant water table below, developing a dawn leafwatêr potential of --0.25 MPa. Water stresses equivalentto -0.7 or -1.0 MPa were developed over 14 d, causing reducedstem and bud elongation, leaf expansion, and bud differentiationand survival. Apex size, the proportion of buds which were floralor vegetative, the early phases of floral initiation, and seedformation on advanced inflorescences were little affected duringthe water deficit period. Upon rewatering previously stressed plants showed increasesrelative to control plants in the rates of shoot appearance,leaf expansion and new node appearance. The ratio of buds becomingfloral was independent of watering treatment, and the enhancedrate of floral bud production in the previously-stressed treatmentswas due to higher rates of total bud differentiation which persistedfor up to six weeks after rewatering. Survival of floral budswas reduced by previous stress, but number of flowers per inflorescence,pod setting, seed number per pod and 100-seed weight were independentof treatment. Seed production was controlled by inflorescencedensity. Rate of seed production was independent of treatmentduring water deficit and four weeks subsequently, and was thenenhanced by 46 and 54 per cent relative to the control in the–0.7 and –1.0 MPa treatments respectively. Macroptilium atropurpureum, Siratro, floral initiation, flowering, seed production, water stress, bud development     

Water Relations of White Clover (Trifolium repens): Water Potential Gradients and Plant Morphology

Water potential, osmotic potential, pressure potential and relativewater content were measured in stolons and leaves of white cloverplants grown under a range of conditions of water supply andevaporative demand. The importance of adventitious roots fromthe nodes was examined. Gradients along stolons were alwaysextremely small, of the order of only 01 MPa. Stolon up waterpotential was representative of plant water status regardlessof stolon length, presence/absence of nodal roots, degree ofwater stress and evaporative demand. It is concluded that waterconduction along stolons was very good. Gradients were foundto exist along petioles; they may have a greater resistanceto water flow than stolons. The relationship between water fluxand stem anatomy, and the importance of differential flow ratesthrough stolons and petioles to plant behaviour during waterstress, are discussed. Trifolium repens L., white clover, water relations     

Water Deficit and Inflorescence Development in Zea mays L.

A water deficit imposed during the period of terminal male inflorescenceinitiation and early development reduced both the growth rateand the mature size of that organ in Zea mays (cv. Iochief).Growth and development of the axillary shoots, the potentialfemale inflorescences, was inhibited during the episode of waterdeficit but promoted thereafter. As a result, plants which hadbeen subjected to a water deficit at that period produced 2–3mature cobs and relatively large axillary shoots at the lowernodes, whereas plants supplied with water throughout produceda single mature cob and relatively small axillary shoots. A water deficit imposed during other growth phases did not producethis response and, moreover, a further period of deficit imposedlater in development, following a deficit at the sensitive stage,inhibited the enlargement of the axillary shoots invoked bythe earlier deficit. It did not, however, inhibit the enhancedfloral development of those axillary shoots nor reverse theinhibition of tassel growth. The data are discussed in relation to correlative inhibitionin Zea mays.     

Effect of Sodium Chloride Stress and Nitrogen Source on Respiration, Growth and Photosynthesis in Lucerne (Medicago sativa L.)

The effects of salinity at different light intensities on freshweight growth and on carbon dioxide influx and efflux were examinedin young plants of lucerne (Medicago sativa L.) that had beengrown in solution culture with nitrogen supplied either as nitrateor by a symbiotic rhizobium. Although the inoculated plantsgrew more slowly than those supplied with nitrate, NaCl at alevel equivalent to an osmotic stress of –0.3 MPa didnot reduce the growth rate of either type of plant under a 12h day-length in a growth chamber. With a day-length of 5 h saltstress (0.0 to –0.6 MPa) did not greatly affect grossphotosynthesis of plants grown on nitrate but respiration ratereached a maximum at –0.3 MPa and declined at larger saltconcentrations. Salt diminished both gross and net photosynthesisin the inoculated plants at a 5 h day-length without stimulatingrespiration. The relationship between photosynthesis and respiration as thephoton flux density was successively decreased was used to inferthe effect of salt on maintenance respiration of the plantssupplied with nitrate. Growth and maintenance components ofrespiration could not clearly be separated in the inoculatedplants suggesting that these were unable to maintain themselvesunder the combined stresses of salt and low light intensity.This view was supported by chemical analysis of the plant material.We conclude that the failure of the inoculated plants to adaptto these conditions could be attributed to the greater demandfor assimilates by the rhizobium. Key words: Medicago sativa, Nitrogen source, Salt stress     

Water Stress Effects on Nitrogen Assimilation and Growth of Trifolium subterraneum L. Using Dinitrogen or Ammonium Nitrate

The relative effects of water stress on growth parameters of subterranean clover (Trifolium subterraneum L. cv. Woogenellup) dependent on either N₂ or 8 millimolar NH₄NO₃ for N were examined. Whole-plant carbon exchange rate (CER), acetylene reduction (AR), dry matter production, and Kjeldahl N accumulation were measured on uniform, intact swards of clover that were maintained under adequately watered conditions or were subjected to three cycles of water stress (leaf water potential ≤−30 bar) over an 18-day period. In the absence or presence of water stress, growth rate, net N accumulation rate, and total N concentration of plants dependent on N₂ were 25 to 26, 45 to 50, and 20 to 21% less, respectively, than plants supplied with 8 millimolar NH₄NO₃. The water stress treatment produced less than a 50% decrease in CER regardless of plant N source, a 90% inhibition of AR in plants dependent on N₂, and a 41% decline in dry matter production on both N sources. Water stress decreased reduced N accumulation 55% in N₂-dependent plants and 50% in NH₄NO₃-dependent plants. Changes in growth and N accumulation caused a 10 to 11% decrease in total plant N concentration of water-stressed plants compared to adequately irrigated controls, but water stress decreased the N concentration of tissue synthesized during the 18-day treatment period in N₂-grown plants more than in plants supplied 8 millimolar NH₄NO₃. Thus, the relative effect of water stress on growth under the two N regimes was similar, but N accumulation by N₂-dependent clover was inhibited to a slightly greater extent (P ≤ 0.001) than in NH₄NO₃-dependent plants.     

The Effects of Nitrogen Supply on the Absorption and Distribution of Copper in Red Clover (Trifolium pratense L.) Grown in Flowing Solution Culture with a Low, Maintained Concentration of Copper

The absorption and distribution of Cu in red clover (Trifoliumpratense L.) were measured in plants grown in flowing solutionculture with Cu maintained throughout at 0.5 µg 1^–1and N supplied either as nitrate or through symbiotic fixation.Although there was a decrease in Cu absorption, both with time,and with a depleted nitrate supply, it increased to its formerrate when nitrate was adjusted to 10 mg N 1^–1 after aperiod of depletion. Differences in absorption between plantsdependent upon fixation and those supplied with nitrate wererelated to the slower initial growth of the plants fixing N.Considerable proportions (> 30 per cent) of the absorbedCu were retained by the roots. At the final harvest, and withthe exception of plants grown with nitrate adjusted to 0.1 mgN 1^–1 after a period of depletion, the proportion of theCu retained was related to the concentration of N in the roots.The different N treatments produced differences in Cu concentrationin the shoots, and the effects were greater in the youngestfully expanded leaves than in older leaves. Trifolium pratense L., red clover, absorption, copper, flowing solution culture, nitrogen     

Root Growth and Water Uptake by Maize Plants in Drying Soil

Sharp, R. E and Da vies, W. J. 1985. Root growth and water uptakeby maize plants in drying soil.— J. exp. Bot. 36: 1441–1456. The influence of soil drying on maize (Zea mays L.) root distributionand use of soil water was examined using plants growing in thegreenhouse in soil columns. The roots of plants which were wateredwell throughout the 18 d experimental period penetrated thesoil profile to a depth of 60 cm while the greatest percentageof total root length was between 20–40 cm. High soil waterdepletion rates corresponded with these high root densities.Withholding water greatly restricted root proliferation in theupper part of the profile, but resulted in deeper penetrationand higher soil water depletion rates at depth, compared withthe well watered columns. The deep roots of the unwatered plantsexhibited very high soil water depletion rates per unit rootlength. Key words: Maize, roots, water deficit, soil water depletion     

Water deficit and plant competition effects on growth and water-use efficiency of white clover ( Trifolium repens,L.) and ryegrass (Lolium perenne,L.)

The combined effects of soil water deficit and above and below ground interspecific plant competition on the growth, water-use efficiency (WUE), and measured carbon isotopic composition (δ¹³C) values of white clover and ryegrass were studied. White clover and ryegrass were grown in specially designed crates 1) individually; 2) in shoot competition; or 3) in shoot + root competition and either well-watered or at a moderate or severe soil water deficit. The effects of shoot + root competition on shoot dry matter growth were substantial and benefited both white clover and ryegrass when well-watered or at a moderate soil water deficit, while severely reducing white clover shoot dry matter growth at severe soil water deficit. Plant competition did not affect the WUE of white clover or ryegrass. As soil water deficit increased, the WUE of white clover did not change whereas the WUE of ryegrass increased and was greater than that of white clover. This was attributed to the lower leaf water conductance of ryegrass which conserved water and maintained growth longer compared to white clover. A stronger correlation existed between soil water deficit and measured δ¹³C values for ryegrass at each plant competition level (P<0.001) than existed for white clover (individual: P<0.01; shoot + root: P<0.001; shoot: P<0.10). Unlike white clover, the relationship between measured δ¹³C values and shoot dry matter growth indicated that C assimilation for ryegrass was dependent on type of plant competition. That WUE remained constant for white clover while measured δ¹³C values increased as soil water deficit increased, suggests that the role below ground respiration rate played in determining δ¹³C values increased. The WUE of white clover appears to be independent of the nature of the competition between plants and the soil water deficit level at which it is grown, whereas for ryegrass, the addition of root competition to shoot competition should lead to increases in its WUE. This revised version was published online in June 2006 with corrections to the Cover Date.     

Induction and Repression of CAM in Sedum telephium L. in Response to Photoperiod and Water Stress

Lee, H. S. J. and Griffiths, H. 1987. Induction and repressionof CAM in Sedurn relephluni L. in response to photopcnod andwater stress.—J. exp. Bot. 38: 834–841. The introduction and repression of CAM in Sedurn telephiunmL, a temperate succulent, was investigated in watered, progressivelydrouglited and rewatered plants in growth chambers. Measurementswere made of water vapour and CO₂ exchange, titratable acidity(TA) and xylem sap tension. Effects of photoperiod were alsostudied. CAM was induced by drought under long or short days,although when watered no CAM activity was expressed. C₃-CAM intermediate plants were used for the investigation ofwater supply. Those which had received water and those drought-stressedboth displayed a similar nocturnal increase in TA, with a day-nightmaximum (H+) of 69 µmol g^–1 fr. wt. The wateredplants took up CO₂ at a maximum rate of 2?2 µmol m^–2s^–1 only in the light period, while the droughted plantsshowed a maximum nocturnal CO₂ uptake rate of 0?69 µmolm^–2 s^–1. Subsequently, as CAM was repressed, thewatered S. telephiwn displayed little variation in TA, withconstant levels at 42 µmol g^–1 fr. wt. (day 10).After 10 d of drought stress, the CAM characteristics of S.telephiurn were aLso affected, with reduced net CO₂ uptake andH+. The transition between C₃ and CAM in S. telephium can be describedas a progression in terms of the proportion of respiratory CO₂which is recycled and refixed at night as malic acid, in comparisonwith net CO₂ uptake. Recycling increased from 20% (day 1) to44% (day 10) as a result of the drought stress and was highin both the CAM-C₃ stage (no net CO2 uptake at night) and alsoin the drought-stressed CAM stage (reduced net CO₂ uptake atnight). The complete C₃-CAM transition occurred in less than8 d, and the stages could be characterized by xylem sap tensionmeasurements: CAM = 0?50 MPa C₃-CAM = 0?36 MPa C₃ = 0?29 MPa. Key words: CAM, Sedum telephium L., recycling     

Osmotic Adjustment and Stomatal Response to Water Deficits in Maize

A pot experiment was carried out using five maize {Zea maysL.) cultivars under three soil moisture levels (MPa 0 to –0.05,–0.3 to –0.9 and –1.2 to –1.5) to investigatethe effects of water deficits on osmotic adjustment and stomatalconductance. The degree of leaf rolling and the sugar and nutrientconcentrations in leaf cell sap were measured. Leaf water potential and osmotic potential decreased and stomatalconductance decreased with increasing water deficits. Stomatalconductance correlated positively with leaf water potentialand osmotic potential. Degree of leaf rolling was lower in cultivarswhich maintained higher turgor. Osmotic adjustment of 0.08 to0.43 MPa was found under the lowest soil moisture level in fivecultivars used. Sugar and K were the major osmotic substancesin the maize plant. Sugar, K and Mg concentrations increasedunder water deficit, and correlated negatively with a decreasein osmotic potential. Key words: Zea mays L., leaf water relations, leaf rolling, osmotic adjustment, stomatal conductance, water deficit     

SHORT COMMUNICATION Growth and Development of White Clover (Trifolium repensL.) as Influenced by P and K Nutrition

The effects of varying P and K supplies on the growth and morphologyof white clover were investigated in a pot experiment. Plantswere treated with three rates of potassium (K) and four ratesof phosphorus (P) in factorial combination, and five harvestswere taken between August 1993 and February 1994. During establishment,shoot and leaf development were more adversely affected by Pthan by K deficiency but, by the final growth period, K deficiencywas the more limiting to growth, causing dramatic reductionsin leaf area index (LAI), numbers of growing points, and stolonand root masses. The adverse affects of K deficiency on shootdevelopment were attributed to the curtailment of petiole andlamina expansion which have major requirements for K. It wasnoted that, under K deficiency, proportionately less assimilatewas partitioned to leaf production than to stolon and root growth;this may be an adaptation enabling K-deficient plants to exploitnutrient supplies in adjacent localities. The results suggestthat, provided white clover can withstand ‘moderate’P stress during establishment, its persistence in swards willprobably be limited more by K deficiency than by low or inadequateP supplies. However, because the experiment was conducted outsidethe normal growing season and with only a single white clovercultivar, it is inappropriate to make any generalizations regardingthe effects of P and K nutrition on white clover in the field.Copyright1998 Annals of Botany Company Mineral nutrition, morphology, phosphorus, potassium,Trifolium repensL.     

Influence of Endophyte and Water Regime Upon Tall Fescue Accessions. I. Growth Characteristics

Superior growth and persistence has been reported in endophyte-infectedgrasses; however, findings may have been confounded by experimentconditions including plant genotype. A controUed-environmentstudy was designed to address some growth characteristics offour tall fescue (Festuca arundinacea Schreb.) accessions asinfluenced by endophyte (Acremonium coenophiahim Morgan-Joneset Gams) and water regime. Endophyte-infected plants were collected,vegetatively propagated and some treated with propiconazole(11 kg a.i. ha^–1) to develop non-infected isolines ofeach accession. The phenotypically diverse accessions, eachrepresented by infected and non-infected isolines, were grownwith adequate (–0–03 MPa), or a series of deficit(<–1·5 MPa) and recovery water regimes, replicatedthree times. Plant growth characteristics were measured during(leaf elongation and tillering) and upon conclusion (phytomassproduction, tillering, and leaf area) of the study. Leaf elongation,as a function of leaf length, was significantly different amongaccessions, and generally decreased with water deficit althoughsome non-infected isolines were not affected. Water deficitdepressed tiller production in virtually all accessions whileendophyte effects depended upon accession. Leaf blade yieldwas not significantly influenced by endophyte status or interactionof endophyte, with water regime and accession; however, pseudostem(stem base and leaf sheath), root and dead leaf yields wereaffected in some cases. Non-structural carbohydrate concentrationin all plant parts except roots, was decreased by water deficit,whereas root non-structural carbohydrate concentration tendedto increase with water deficit. Non-structural carbohydratesof all plant parts was not influenced by endophyte status. Tallfescue-endophyte association responses vary due to genotype,therefore a simple generalization of endophyte impact upon tallfescue productivity and persistence is not possible based uponthe results of this study Festuca arundinacea Schreb., Acremonium coenophiahim Morgan-Jones et Gams, leaf elongation, phytomass production, tillering, water deficit, non-structural carbohydrate     

Water deficit-induced oxidative stress and the activation of antioxidant enzymes in white clover leaves

The objective of this study was to determine the development of the antioxidant enzymes induced by drought stress in white clover (Trifolium repens L.) leaves. Water stress was imposed during 28 d by decreasing the daily irrigation. Leaf water potential (Φ_w) gradually decreased from −0.46 to −2.33 MPa. For the first 7 d, dry mass (DM), H₂O₂ and lipid peroxidation were not significantly affected by water deficit. From 14 d of treatment, water stress decreased dry mass and increased content of reactive oxygen species (O₂ ^·− and H₂O₂) and oxidative stress (malondialdehyde content). The ascorbate peroxidase (APOD) was activated most rapidly, already during the first week of water stress, but then its activity slowly decreased. Activation of superoxide dismutase (SOD) and catalase (CAT) by water deficit continued during the 14 d (Φ_w ≥ −1.65 MPa) and then their activities remain on the similar level. The activity of guaiacol-peroxidase (GPOD) increased mostly under progressive water stress and was correlated with increase in lipid peroxidation and growth restriction.     

The Effect of Water Stress upon Polyamine Levels in Barley (Hordeum vulgare L.) Leaves

The effect of water stress on leaf polyamine content of fourHordeum vulgare varieties, Alger/Ceres, Palmella Blue, Rihaneand Roho, with different drought characters was studied After6 d without water Alger/Ceres, Palmella Blue and Rihane hadaccumulated putrescine, although only in concentrations up totwice those found in the controls, but Roho had a decreasedputrescine content. However, one common response was identified;the accumulation and subsequent loss of putrescine was dependentupon the maintenance and loss of leaf turgor respectively. Consequentlyvarietal differences in putrescine accumulation were relatedto water consumption rates and the extent of osmotic adjustment.Spermine behaved in a similar manner to putrescine but spermidinelevels always decreased. Polyamine levels were never high enoughto be an important component of solute accumulation. Prolinelevels were 150-fold higher and glycine-betaine levels 50-foldhigher than polyamines in stressed plants. Proline and glycine-betaineaccumulation occurred once a threshold turgor was reached, whichin Roho appeared to be a reduction by 0•2–0•25MPa or 30–40%. The importance of polyamine accumulationduring water stress is discussed. Key words: Hordeum vulgare, polyamine, water stress     

Apical Dominance, Water Deficit and Axillary Inflorescence Growth in Zea mays: The Role of Abscisic Acid

In the sweet corn cultivar, Iochief, an episode of water deficitduring early tassel development results in a subsequent promotionof the growth of the lower axillary inflorescences. This responseis also produced by the application of abscisic acid (ABA) atthis period of growth to well-watered plants, and the hypothesisthat the response to water deficit was due to an increase inendogenous ABA concentration was examined. The ABA contentsof the tassel, leaf and axillary inflorescences were found toincrease during water stress, the increase in the tassel andaxillary buds being most rapid in the first 2 days of waterdeficit. This increase in free ABA content was followed after4 days of water deficit by a progressive increase in the concentrationof ‘bound’ ABA in the tissues. There was littleincrease in free ABA concentration after 4 days water deficit;this paralleled the subsequent growth response of the axillaryinflonscences which also was unaffected by prolonging the epidoseof water deficit beyond 4 days. In order to establish whether the response of the axillary inflorescencesto ABA was dependent upon the presence of the tassel, ABA wasapplied to watered plants with or without the developing tassel.As had been previously found with water stress, removing thetassel inhibited the response of the plant to applied ABA. Zea mays, apical dominance, water stress, inflorescence growth, abscisic acid     

Some Effects of Abscisic Acid and Water Stress on Stomata of Vicia faba L.

Vicia faba seedlings grown under a plastic tent in the laboratorywere either watered well throughout their growth period or weresubjected to a water stress treatment for several days priorto an experimental treatment. The effects of a further waterstress treatment or an application of an aqueous solution ofabscisic acid (ABA) on the stomata of these plants were determined.Stomata of previously water-stressed plants proved to be moresensitive than stomata of well watered plants to ABA appliedthrough the petiole via the transpiration stream and sprayedonto leaf surfaces. Stomata of previously water-stressed plantsclosed more rapidly and to a greater degree than stomata ofwell watered plants. The hormone had only a small effect whenapplied directly to epidermal fragments removed from both groupsof plants. Stomata of plants which had received a water stresspretreatment were less sensitive to a subsequent period of waterstress than were stomata of previously well watered plants.It is proposed that stomatal adaptation to water stress maybe related to changes in the hormonal balance of the plant.     

The Responses of Leaf Water Potential and Leaf Diffusive Resistance to Abscisic Acid, Water Stress and Low Temperature in Hibiscus esculentus: The Effect of Water Stress and ABA Pre-Treatments

Intact plants of okra (Hibiscus esculentus) were chilled at6°C in the light, and leaf diffusion resistance (LDR) andleaf water potential measured. The response of the LDR of excisedleaves to fresh weight loss and, separately, exogenous abscisicacid (ABA) supply, was also studied at 6°C and 30°C.The influence of two pre-treatments upon these measurementswas studied. The two pre-treatments consisted of the impositionof a period of water stress at 30°C prior to measurement(followed by re-watering) or the spraying of the leaves withABA. It was found that plants that had been grown in a highhumidity environment continuously (designated control plants)had stomata that were very unresponsive to both water loss fromthe leaves and to exogenous ABA at both temperatures (6°Cand 30°C). Chilling the control plants resulted in rapidwilting and concomitant decline in leaf water potential. A pre-treatmentof water stress prior to chilling did not alter or reduce therate of development of chilling injury, nor did the pre-treatmentincrease the responsiveness of stomata at 6°C to water lossor exogenous ABA. However, spraying the leaves with ABA priorto chilling reduced the severity and delayed the onset of chillinginjury. Stomatal response to water stress and exogenous ABAwas increased by the spraying pre-treatment. These results arediscussed in relation to previous studies of the phenomenonof stomatal locking open at low temperature and the effect ofpre-treatments upon the development of chill-resistance. Key words: Water stress, chilling, stomata     

Effects of Blue Light on the Vertical Colonization of Space by White Clover and their Consequences for Dry Matter Distribution

The morphology of white clover is very sensitive to the lightenvironment, especially to the ratio of red:far-red light andto photon irradiance. However, less is known about the effectsof blue light on clover morphogenesis. Cuttings of white cloverwere grown for 56 d in two controlled chambers receiving lightwith similar photosynthetic efficiency and phytochrome photoequilibriumstate but different levels of blue light: some plants were grownunder orange light (very low blue light, 0.02 µmol m^-2s^-1)or under white light containing blue light (83 µmol m^-2s^-1).Other plants were switched from white light to orange lightorvice versa,after 30 d. The absence of blue light modifiedthe growth habit of clover and raised the laminae in the upperlayer of the canopy by increasing petiole length, and petioleangle from the horizontal, and by raising stolons above theground surface. Moreover, the absence of blue light had no effecton total leaf area and total dry weight per plant, but increasedthe leaf area and biomass of petioles of the main axis. Largerpetioles and laminae were associated with the allocation ofmore dry weight to the petiole at the same petiole thicknessbut with thinner laminae. These results indicate that a decreasein blue light is involved in the perception of, and adaptationto, shading by the plant.Copyright 1997 Annals of Botany Company Biomass allocation; blue light; growth habit; leaf area; light quality; photomorphogenesis; Trifolium repensL.; white clover