Influence of Endophyte and Water Regime Upon Tall Fescue Accessions. II. Pyrrolizidine and Ergopeptine Alkaloids

Alkaloids, along with specific environmental conditions, havebeen associated with both detrimental and beneficial aspectsof endophyte (Acremonium coenophialum Morgan-Jones et Gams)infected tall fescue (Festuca arundinacea Schreb.) associations.Benefits to the plant accrue through reduced herbivory, whereasdetriment to the animal occurs as altered grazing behaviourand reduced productivity. A controlled environment study wasconducted to examine pyrrolizidine and ergopeptine alkaloidconcentration of four tall fescue accessions as influenced byendophyte status and water regime. Endophyte-free plants weredevoid of ergopeptine alkaloid and contained little, if any,pyrrolizidine alkaloid. Leaf blade tissue of endophyte-infectedisolines contained a range of both ergopeptine (256 to 1633ng g^–1) and pyrrolizidine (92 to 450 µg g^–1)alkaloid concentrations. Water deficit generally increased alkaloidconcentration. Alkaloid yield, based upon concentration andtissue d. wt, showed that significant increase in ergopeptineand pyrrolizidine alkaloid in leaf tissue was associated withwater deficit and was due to actual increased synthesis andnot simply decreased phytomass. Leaf and pseudostem (leaf sheathand stem base) tissue alkaloid concentrations indicated differentaccumulation patterns for ergopeptine and pyrrolizidine alkaloids.Ergopeptine alkaloid yield increased in water-stressed pseudostem,whereas pyrrolizidine alkaloid yield decreased in some, butnot all accessions. The range of host genotype/endophyte biotyperesponse offers the possibility to select associations whichproduce few deleterious effects in animals yet maintain highforage productivity and persistence. Festuca arundinacea, Acremonium coenophialum, tall fescue genotypes, water stress, N-formyl and N-acetyl loline, ergovaline     

Does Endophyte Influence Regrowth of Tall Fescue?

The influence of ecological and environmental factors on theresponse of the mutualistic symbionts Acremonium coenophialum(Morgan-Jones and Gams), and tall fescue ( Festuca arundinacea,Schreb.) has received substantial attention. However, much ofwhat is known about developmental and physiological responseshas been obtained from static or mature canopies and the influenceof defoliation on the symbiota has not been considered in detail.A controlled environment experiment was conducted to assessthe influence of defoliation and endophyte on the regrowth anddevelopment of two tall fescue genotypes and their respectiveendophytes, known to differ in morphology and alkaloid productioncapacity. The response of endophyte infected relative to non-infectedplants was evaluated for measured and calculated parametersfor each defoliation. While the influence of genotype was substantialon virtually all parameters, endophyte interacted with genotypeand defoliation resulting in enhanced plasticity of mutualistsas a response to varying conditions. Endophyte infection influencedleaf mass depending upon genotype, while the relative benefitof endophyte on pseudostem mass was affected by defoliation.Endophyte interacted with genotype to influence relative growthrate and productivity relative to nitrogen concentration whencanopies were clipped, but was less important when canopiesremained uncut. In some instances endophyte gave growth andsize advantage to the host and did not in others. Generalitiesregarding symbiont response to extrinsic factors may be misleading,since responses depend upon the specific mutualism consideredand conditions imposed. Acremonium coenophialum ; Festuca arundinacea ; non-structural carbohydrate; relative growth rate     

Estimation of Potential Tiller Production and Site Usage During Tall Fescue Canopy Development

Tiller appearance in tall fescue (Festuca arundinacea Schreb.)occurs in an orderly, predictable manner with the potentialfor a high degree of synchronization among tillers on a givenplant. Estimates of potential cumulative tiller production (Tmax)are made for synchronous (Tmax_lx = 2^Lx+1 – 1, where Lxis the axil number of the youngest leaf on the main stem whichbears an emerged tiller) and non-synchronous (Tmax₁ = 2^L–NLAT–1,where L is the number of leaves on the main stem and NLAT thenumber of leaves above the youngest primary tiller at its appearance)conditions. A method for determining the degree of synchronizationand an equation for estimating site usage are also presented.Early in seedling development, site usage of a tall fescue populationwith high tillering capacity was near 90%, and tillering wasregulated largely by rate of tiller site formation. As the canopydeveloped the phyllochron (time between successive leaf appearances)and NLAT increased, slowing the rate of tiller production intemporal terms and in relation to leaf appearance, respectively.Beyond 45 d after planting, site usage decreased rapidly, furtherreducing tiller production. High tiller production appearedto be associated with synchronized tiller appearance, with alack of synchrony being associated with decreased site usage.Tillers formed in prophyll axils were less likely to be in synchronywith other tillers and frequently failed to appear. In contrastwith branching in dicotyledons, apical dominance appears toplay a minor role in regulating tillering in tall fescue Tillerproduction initially appears to proceed at near maximum ratesthen is down-regulated during later development by longer phyllochrons,slower rate of tiller elongation and reduced site usage. Festuca arundinaceaSchreb., tall fescue, tiller production, leaf appearance, site usage, leaf elongation rate, synchronization, phyllochron, canopy development     

Defoliation and Leaf Age Influence on Ergot Alkaloids in Tall Fescue

A controlled environment experiment was conducted to determinethe influence of defoliation on the regrowth and developmentof two tall fescue (Festuca arundinacea Schreb.) host-endophyte(Neotyphodium coenophialum Morgan-Jones and Gams (Glenn, Bacon,Price and Hanlincomb. nov. ) associations (DN2 and DN11), knownto differ in morphology and alkaloid production capacity. Defoliationtreatments included an uncut control, and clipping to a 5- or10-cm residue height. In a separate experiment, leaf age effectswere determined on an uncut plant canopy. Ergot alkaloid concentrationwas greatest in pseudostem and least in harvested leaf. Non-infectedplants were devoid of alkaloid. Alkaloid production (expressedas a function of dry matter) and yield were greater in uncutthan clipped plants, and were greater in DN11 than DN2 plants.Alkaloid production increased with increasing N concentrationin both associations; however, the rate of production was influencedby concentration of non-structural carbohydrate. Leaf age influencedleaf mass and alkaloid concentration and yield. Oldest leaves(>6 weeks after appearance) of DN2 and DN11 had the lowestconcentrations of ergot alkaloid, while leaves that were 2 to4 weeks old had the greatest alkaloid concentration and yield(alkaloid concentrationxleaf mass). Repeated defoliation reducedthe production of alkaloids, but did so as a function of non-structuralcarbohydrate. Our data suggest that alkaloid production canbe modified by defoliation as well as by host-endophyte association. Neotyphodium coenophialum ; Festuca arundinacea ; N; non-structural carbohydrate     

Nitrogen Effects on Leaf Anatomy within the Intercalary Meristems of Tall Fescue Leaf Blades

Longitudinal elongation contributes most to leaf area expansionof grasses and its rate is known to be strongly affected byN. Our objective was to determine the effect of two N regimes(N₀and N+) on the gradient of leaf tissue formation in meristemsof two contrasting tall fescue (Festuca arundinacea Schreb.)genotypes. Proportions of epidermal, mesophyll and vasculartissue as well as intercellular air space were determined throughoutthe base of actively elongating leaves. The area of leaf transversesections nearly doubled between the ligule and the distal endof the growth zone (about 30 mm), and was mainly associatedwith lateral epidermal and mesophyll cell division in the proximal5.0–7.5 mm. Further increase in transverse area was dueto the formation of intercellular airspace and transverse expansionof epidermal cells. Depending on genotype and N treatment themesophyll, epidermis, vascular bundles and air space comprised45–54%, 20–28%, 6–9%, and 17–21%, respectively,of transverse leaf area in the distal part of the growth zone.After a slight increase close to the leaf base, the area ofvascular tissue remained constant throughout the growth zone.The proportion of air space to mesophyll space was higher atN₀than at N+ because mesophyll area was enhanced by N+ to agreater degree than by N₀. In the genotype with slow leaf elongation,the increase in cross-sectional leaf area was due to an increasein both leaf width and leaf thickness. In the genotype whichhad faster leaf elongation and wider leaves, only leaf thicknesswas enhanced by N+. Copyright 2001 Annals of Botany Company Festuca arundinacea(Schreb.), tall fescue, leaf anatomy, growth zone, nitrogen     

Epidermal Cell Division and the Coordination of Leaf and Tiller Development

Initiation and development of grass leaves and tillers are oftendescribed individually with little attention to possible interrelationshipsamong organs. In order to better understand these interrelationships,this research examined epidermal cell division during developmentaltransitions at the apical meristem of tall fescue (Festuca arundinaceaSchreb.). Ten seedlings were harvested each day for a 9-d period,and lengths of main shoot leaves and primary tillers were measured.In addition, numbers and lengths of epidermal cells were determinedfor 0·5 mm segments along the basal 3 mm of each leafand tiller. Primordia development and onset of rapid leaf elongationwere characterized by an increase in the number of cells perepidermal file with mean cell length remaining near 20 µmper cell. After the leaf had lengthened to 1-1·5 mm,cells near the leaf tip ceased dividing and increased in length,at which time leaf elongation rate increased rapidly. Liguleformation, marking the boundary between blade and sheath cells,occurred prior to leaf tip emergence above the whorl of oldersheaths, while the earliest differentiation between blade andsheath cells probably began when leaves were < 1 mm long.Major transitions in leaf and tiller development appeared tobe synchronized among at least three adjacent nodes. At theoldest node, cessation of cell division in the leaf sheath wasaccompanied by initiation of cell division and elongation inthe associated tiller bud. At the next younger node the ligulewas being initiated, while at the youngest node cell divisioncommenced in the leaf primordium, as elongation of a new leafblade began. This synchronization of events suggests a key rolefor the cell division process in regulating leaf and tillerdevelopment.Copyright 1994, 1999 Academic Press Festuca arundinacea Schreb., tall fescue, cell division, leaf initiation, tillering, ligule development     

Specific Leaf Weight in Zones of Cell Division, Elongation and Maturation in Tall Fescue Leaf Blades

Patterns of change in specific leaf weight (SLW), water-solublecarbohydrate (WSC) content and leaf width were used to delineatethe region of secondary cell wall accumulation, and determinethe rate of increase in structural material along a developingleaf blade of tall fescue (Festuca arundinacea Schreb.). Structuralspecific leaf weight (SSLW) was determined by subtracting WSCmass from dry weight to emphasize structural material. Becausemeristematic activity, cell elongation, and cellular maturationare arranged successively in the grass leaf, these patternsrepresent a developmental sequence through which each segmentof the leaf blade passes. Patterns were generally similar fortwo genotypes, one selected for high (HYT) and the other forlow (LYT) yield per tiller, for a single genotype grown at 17or 25 C, and for two field-grown populations which differedin leaf area expansion rate (LAER). In all three studies, the elongation zone of the developingleaf had 31 to 39 per cent WSC on a dry weight basis. The LYTgenotype had a higher SLW at all stages of development whengrown at 17 than at 25 C, due to greater WSC accumulation.At 20 C, the HYT genotype had a higher SLW all along the elongatingleaf blade than the LYT genotype. This difference was due toa difference in SSLW, while WSC content was similar. The LERwas 64 per cent higher in the high population than the low,but elongation zones were similar in WSC. In all cases, SSLWwas high in the meristematic region, lowest near the distalend of the cell elongation zone, then increased linearly astissue matured. Rate of increase in SSLW was 8.5 and 5.2 g m^–2d^–1 for the HYT and LYT genotypes, respectively, and 7.6and 6.7 g m^–2 d^–1 for the high and low LAER populations,respectively. Festuca arundinacea Schreb., tall fescue, specific leaf weight, leaf width, water-soluble carbohydrates, leaf elongation rate     

The Effect of Wind on Grasses: 1. CUTICULAR AND STOMATAL TRANSPIRATION

Transpiration of Festuca arundinacea Schreb, strain S170, wasmeasured at two different wind speeds in a controlled-environmentwind tunnel. As a result of a wind increase from 1 m s^–1to 3.5 m s^–1 above the sward, transpiration graduallyincreased, especially at night-time. Similar effects were notedin three other grass species. The transpiration increase couldbe attributed to decreases in stomatal and cuticular resistances,as a result of leaf buffeting.     

The effect ofAcremonium coenophialum on the growth and nematode infestation of tall fescue

The presence of the endophytic fungusAcremonium coenophialum Morgan-Jones et Gams in tall fescue (Festuca arundinacea Schreb.) induces toxicity when this grass is grazed by cattle; however, there is evidence that removing the endophyte reduces the stand vigor and longevity of fescue. A field trial was conducted to determine the effects of water supply and the presence of the endophytic fungus on plant growth, drought tolerance, and soil nematode populations in Kentucky 31 tall fescue. The design included two factors, level of endophyte infection (0 and 75%) and irrigation regime (none, low, and high). Where water deficits occurred, herbage yield and leaf area were lower, and percentage dead tissue and canopy minus air temperature were greater in endophyte-free compared with endophyte-infected fescue. Soil populations ofPratylenchus scribneri andTylenchorhynchus acutus were substantially higher in the noninfected than in the endophyte-infected plots. The endophyte apparently confers drought tolerance to Kentucky 31 tall fescue, and this effect may be at least partially mediated through enhanced resistance to soil-borne nematodes.Published with the approval of the Director of the Ark. Agric. Exp. Stn.     

An Ecophysiological Comparison of Measurements of the Diurnal Rhythm of the Leaf Elongation and Changes of the Leaf Thickness of Salt-resistant Dicotyledonae and Monocotyledonae

Rozema, J., Arp, W., van Diggelen, J., Kok, E. and Letschert,J. 1987. An ecophysiological comparison of measurements of thediurnal rhythm of the leaf elongation and changes of the leafthickness of salt-resistant Dicotyledonae and Monocotyledonae.—J.exp. Bot. 38: 442–453. The continuous measurement of leaf elongation and leaf thicknesswith the use of a rotation potentiometer set up revealed a rapidand sensitive reaction of halophytic plants to conditions affectingthe plant's water relations. At increased salinity (450 molm^–3 NaCl) the rate of leaf elongation decreased both inAster tripolium and in Sparlina anghca. Increased shrinkageduring the day and a long period for recovery swelling at nightin leaves of Aster iripolium at increased salinity illustratesthat water shortage is part of the cause of salinity-inducedgrowth reduction. All dicotyledonous species analysed (Aster tripolium, A triplexhastata, A. littoralis, Suaeda maritima and Beta vulgaris) showeda day/night ratio of the leaf elongation rate lower than 1,while this ratio was higher than or equal to 1 in Monocotyledons(Spartina anglica, Juncus gerardii, J. maritimus, Festuca rubrassp. litoralis, Elymus pycnanthus). With the exception of Triglochinmaritima none of the monocotyledonous halophytes tested (Sparlinaanglica, Juncus gerardii, J. maritimus, Festuca rubra ssp. litoralis,Elymus pycnanthus) exhibited a diurnal rhythm of leaf thicknesschanges, such as was observed for all dicotyledonous speciesstudied (Aster tripolium, Atriplex hastata, A. littoralis, Salicorniabrachyslachya, Suaeda maritima, Glaux maritima, Odontites vernassp. serotina). The diurnal pattern of the leaf elongation rateand the leaf thickness changes can be explained by variationof photosynthetic rate and transpiration water losses by stomatalclosure in the dark and opening in the light such as shown forthe dicotyledon species Glaux maritima. This difference betweendicot and monocot species in diurnal variation of the leaf elongationrate and leaf thickness may partly be explained in terms ofthe different position of the growth zone and possibly by adifference in elasticity of the tissue of halophytic monocotyledonsand dicotyledons. The consequences of these differences arediscussed. Key words: Leaf elongation rate, leaf thickness, water relations, salt resistance, Dicotyledonae, Monocotyledonae     

Relationships among non-Acremonium sp. fungal endophytes in five grass species.

Many cool-season grasses (subfamily Pooideae) possess maternally transmitted fungal symbionts which cause no known pathology and often enhance the ecological fitness and biochemical capabilities of the grass hosts. The most commonly described endophytes are the Acremonium section Albo-lanosa spp. (Acremonium endophytes), which are conidial anamorphs (strictly asexual forms) of Epichloë typhina. Other endophytes which have been noted are a Gliocladium-like fungus in perennial ryegrass (Lolium perenne L.) and a Phialophora-like fungus in tall fescue (Festuca arundinacea Schreb.). Here, we report the identification of additional non-Acremonium sp. endophytes (herein designated p-endophytes) in three more grass species: Festuca gigantea, Festuca arizonica, and Festuca pratensis. In each grass species, the p-endophyte was cosymbiotic with an Acremonium endophyte. Serological analysis and sequence determinations of variable portions of their rRNA genes indicated that the two previously identified non-Acremonium endophytes are closely related to each other and to the newly identified p-endophytes. Therefore, the p-endophytes represent a second group of widely distributed grass symbionts.     

Molecular biology and evolution of the grass endophytes.

Acremonium coenophialum Morgan-Jones et W. Gams is a maternally transmitted fungal symbiont (endophyte) of the important forage grass Festuca arundinacea Schreb. (tall fescue), and provides biological protection and enhanced fitness to its host, but its anti-mammalian ergot alkaloids detract from the usefulness of tall fescue as forage for livestock. Molecular genetic techniques and materials are being developed in order to specifically eliminate the gene(s) encoding the first enzyme in ergot alkaloid biosynthesis. These techniques will also facilitate basic studies, such as host-fungus compatibility or biosynthesis of insecticidal alkaloids. Molecular phylogenetics indicate that endophytes related to A. coenophialum have evolved on multiple occasions from strains of Epichlo? typhina (Ascomycotina, Clavicipitaceae), for which the sexual cycle is known. These studies also reveal significant diversity among seedborne endophytes in individual grass species. Thus, the endophytes are an important source of biochemical potential and genetic diversity in grass-fungus symbiota.     

Effects of elevated CO2 concentrations on three montane grass species: III. Source leaf metabolism and whole plant carbon partitioning

Agrostis capillaris L.⁵, Festuca vivipara L. and Poaalpina L.were grown in outdoor open-top chambers at either ambient (340 3µmol mol^–1) or elevated (6804µmol mol^–1)concentrations of atmospheric carbon dioxide (CO₂) for periodsfrom 79–189 d. Photosynthetic capacity of source leaves of plants grown atboth ambient and elevated CO₂ concentrations was measured atsaturating light and 5% CO₂. Dark respiration of leaves wasmeasured using a liquid phase oxygen electrode with the buffersolution in equilibrium with air (21% O₂, 0.034% CO₂). Photo-syntheticcapacity of P. alpina was reduced by growth at 680 µmolmol^–1 CO₂ by 105 d, and that of F. vivipara was reducedat 65 d and 189 d after CO₂ enrichment began, suggesting down-regulationor acclimation. Dark respiration of successive leaf blades ofall three species was unaltered by growth at 680 relative to340 µmol mol^–1 CO₂. In F. vivipara, leaf respirationrate was markedly lower at 189 d than at either 0 d or 65 d,irrespective of growth CO₂ concentration. There was a significantlylower total non-structural carbohydrate (TNC) concentrationin the leaf blades and leaf sheaths of A. capillaris grown at680µmol mol^–1 CO₂. TNC of roots of A. capillariswas unaltered by CO₂ treatment. TNC concentration was increasedin both leaves and sheaths of P. alpina and F. vivipara after105 d and 65 d growth, respectively. A 4-fold increase in thewater-soluble fraction (fructan) in P. alpina and in all carbohydratefractions in F. vivipara accounted for the increased TNC content. In F. vivipara the relationship between leaf photosyn-theticcapacity and leaf carbohydrate concentration was such that therewas a strong positive correlation between photosynthetic capacityand total leaf N concentration (expressed on a per unit structuraldry weight basis), and total nitrogen concentration of successivemature leaves reduced with time. Multiple regression of leafphotosynthetic capacity upon leaf nitrogen and carbohydrateconcentrations further confirmed that leaf photosynthetic capacitywas mainly determined by leaf N concentration. In P. alpina,leaf photosynthetic capacity was mainly determined by leaf CHOconcentration. Thus there is evidence for down-regulation ofphotosynthetic capacity in P. alpina resulting from increasedcarbohydrate accumulation in source leaves. Leaf dark respiration and total N concentration were positivelycorrelated in P. alpina and F. vivipara. Leaf dark respirationand soluble carbohydrate concentration of source leaves werepositively correlated in A. capillaris. Changes in source leafphotosynthetic capacity and carbohydrate concentration of plantsgrown at ambient or elevated CO₂ are discussed in relation toplant growth, nutrient relations and availability of sinks forcarbon. Key words: Elevated CO₂, Climate change, grasses, carbohydrate partitioning, photosynthesis, respiration     

The Effect of Wind and a Reduced Supply of Water on the Growth and Water Relations of Festuca arundinacea Schreb

Festuca arundinacea was grown at high and low wind-speeds attwo levels of soil water. Transpiration was increased at highwind-speed and accompanied by leaf water stress. Growth of leafarea was progressively reduced according to the severity ofthe experimental treatments in the sequence: wet soil and lowwind; dry soil and low wind; wet soil and high wind; dry soiland high wind. The leaf water potential was also reduced inthis sequence. Festuca arundinacea Schreb., transpiration, water stress, wind, water potential     

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     

Drought effects on cellular and spatial parameters of leaf growth in tall fescue

The effect of drought and recovery on cellular and spatial parametersof the growth process in tall fescue leaves was studied in twoexperiments. In both experiments plants grown on vermiculiteand maintained in a controlled environment were submitted toa 7 d drought period generated by withholding water. Droughtwas followed by a 3 d recovery period in experiment II. As leafelongation rate (LER) decreased during developing drought boththe growth zone length (initially 40 mm) and the maximum relativeelemental growth rate (initially 0.09 mm mm^–1 h^–1during the dark period of diurnal cycles) within the growthzone declined. But the growth zone still exhibited a lengthof approximately 15 mm when LER approached 0 under severe drought(–2.0 MPa predawn leaf water potential). The growth potentialof the basal 15-mm-long portion of the leaf was conserved duringthe period when drought effected the complete arrest of leafelongation. A (retrospective) analysis of the position-timerelationships of epidermal cells identified on leaf replicas(experiment II) indicated that the cell flux out of the growthzone responded very sensitively to drought. Before drought theflux was maximum at approximately 3.2 cells (cell file h)^–1during the dark period. Flux decreased to 0 when leaf elongationstopped. Flux also varied diurnally both under well-wateredand droughted conditions. In well-watered conditions it wasabout 30% less during the light than the dark period. Cell elongationwas also sensitive to drought. Under well-watered conditionsepidermal cell elongation stopped when cells attained a lengthof approximately 480 µm. During developing drought cellsstopped elongating at progressively shorter lengths. When LERhad decreased to almost nil, cells stopped elongating at a lengthof approximately 250 µn. When drought was relieved followinga 2 d complete arrest of leaf elongation then cells shorterthan 250 µm were able to resume expansion. Following rewateringcell flux out of the growth zone increased rapidly to and abovethe pre-drought level, but there was only a slow increase overtime in the length at which cell elongation stopped. About 2d elapsed until the leaf growth zone produced cells of similarlength as before drought (i.e. approximately 480 µm). Key words: Epidermal cell length, cell flux, (leaf) growth zone, leaf elongation rate, relative elemental growth rate, position-time relationships (path line, growth trajectory), drought, water deficit     

The Effect of Wind on Grasses: V. LEAF EXTENSION DIFFUSIVE CONDUCTANCE PHOTOSYNTHESIS IN THE WIND TUNNEL

Festuca arundicacea and Lolium perenne were grown in a controlled-environmentwind tunnel at high (7.4 m s^–1) or low (1.0 m s^–1)windspeed. The rate of leaf extension was markedly reduced atthe high windspeed. This effect could not be attributed to waterstress, for, although the leaf conductance increased with exposureto high wind, no effect on leaf water potential was detected.Nor was the rate of photosynthesis affected when the windspeedwas changed. Moreover, concentrations of ethylene in the windtunnel were too low to explain the observations. It is suggestedthat mechanical stimulus itself may have caused the reductionin leaf growth rate.     

Stomatal Control of Water Loss in Siratro (Macroptilium atropurpureum (DC) Urb.), a Tropical Pasture Legume

Stomatal conductance of siratro declined linearly as leaf waterpotential fell until zero conductance was reached at –10bar. In a grass/legume pasture stomata of siratro respondedto humidity (saturation deficit), and to a lesser extent toleaf water potential, such that leaf water potential did notfall below –9 bar, whereas that of the grass continuedto decline for most of the day. The dual response of siratroto both humidity and leaf water potential suggests that thisspecies has an efficient two-stage stomatal control of waterloss which provides an explanation of its higher leaf waterpotential and greater drought avoidance compared with sown grassesin semi-arid areas of north-eastern Australia. Macroptilium atropurpureum (DC) Urb., siratro, Desmodium uncinatum, stomatal control, stomatal conductance, water loss, leaf water potential, drought avoidance, saturation deficit     

Tillering, Leaf Expansion and Growth of Plants of Two Cultivars of Perennial Ryegrass Grown using Hydroponics at Two Water Potentials

Tillering and growth parameters of perennial ryegrass cultivarsWendy (diploid) and Condesa (tetraploid) were determined ina glasshouse experiment using hydroponics at low (–1·3MPa) and normal water potential (0 MPa). At –1·3MPa, leaf extension rate was reduced by 36%. Final plant tillernumber was 20% lower at –1·3 MPa because of a 12%reduction in the leaf appearance rate in the first weeks afterthe start of the treatments. Site filling, the relative increasein tiller number per leaf appearance interval, was high (0.61)-butstill lower than theoretically possible-and was only slightlyaffected by water potential. Site filling was shown to be strictlyrelated to the number of inhibited plus unemerged tiller buds.Dry matter production was 64% lower at –1·3 MPa.Relative growth rate (RGR) was, on average, 17% lower at –1·3MPa, but the reduction was greater just after the treatmentsstarted. Also, net assimilation rate (NAR) was reduced moreby low water potential just after the start of the treatments.Specific leaf area (SLA) was 13 % lower at –1·3MPa for Wendy, but not significantly reduced for Condesa. Contraryto expectations based on the theory of the functional balancebetween root and shoot, leaf weight ratio was slightly higherat –1·3 MPa. From comparison of the results ofthis study with published data, it is concluded that effectsof drought in the field on tillering cannot be attributed onlyto low water potential. Lolium perenne L., perennial ryegrass, tillering, site filling, leaf appearance, leaf extension, growth analysis, water potential