The Effect of (2-chloroethyl)-trimethylammonium Chloride on the Growth of Barley

Barley (Hordeum vulgare L. C.I.666) was shown to be susceptibleto the growth retardant (2-chloroethyl)-trimethylammonium chloride(CCC). The estimation of cell number in the dwarfed third leafblade indicated that a decrease in mitotic activity had occurredin treated plants. There was also a decrease in cell size intreated plants. The dwarfing action of CCC was reversed by exogenousgibberellic acid (GA₃) but this was shown to be the result ofincreased cell elongation only. GA₃ did not promote cell divisionin healthy or CCC-treated plants. Assay of endogenous gibberellinsshowed a significant reduction in the level of a substance correspondingto GA₃ in CCC-treated plants. It is suggested that CCC-induceddwarfing of barley is largely the result of a reduction in meristematicactivity. This may be related to an effect on gibberellin biosynthesisbut is not reversed by the application of exogenous GA ₃.     

The Anther Smut of Sea Campion: A Study of the Role of Growth Regulators in the Dwarfing Symptom

The sympton of dwarfing in the sea campion, Silene vulgaris(Moench) Garcke sub sp. maritima(With.) A. & D. Löveinfected with the anther smut fungus Ustilago violacea (Pers.)Fuckel, a systemic, perennial parasite, has been investigated. Extracts of both healthy and diseased plants contain IAA andGA₃, but diseased plants contain less gibberellins than healthyplants. Neither IAA nor gibberellins were detected in significantquantities in the medium when U. violacea was grown in pureculture, but IAN was present. IAN was also found m extractsof diseased plants and it is tentatively suggested that it isformed by the fungus and may accumulate in the host owing tothe inability of the plant to convert IAN to IAA The exogenousapplication of IAN to healthy plants does not produce any diseasesymptoms. The dwarfing symptom of the diseased plant may be due to thelower levels of gibberellins which it contains compared withhealthy plants since the exogenous application of GA₃, restoresthe diseased plant to normal growth     

The Relationship between Levels of Endogenous Gibberellins and the Response of Vigna Hypocotyls to Exogenous Indole-3-Acetic Acid

Segments excised from the upper and the lower parts of cowpea(Vigna unguiculata L.) hypocotyls were compared in terms oftheir responses to exogenous indole-3-acetic acid (IAA) in relationto their endogenous levels of gibberellin. Growth of the segmentswas measured continuously during xylem perfusion with a lineardifferential transformer. IAA induced a burst of elongationin the upper segments but only slight promotion of growth inthe lower segments. Treatment with uniconazole, a potent inhibitorof the biosynthesis of gibberellins, reduced the responsivenessof the upper segments to exogenous IAA to about one half ofthe control value. Pre-perfusion with GA₃ of such segments fortwo hours prior to application of IAA, partially restored theresponsiveness to IAA. Analysis by GC/MS identified GA₁, GA₄,GA₉ GA₂₀ and GA₅₁ as native gibberellins in the hypocotyls ofcowpea seedlings. Analysis by GC/SIM also showed that the physiologicallyactive gibberellins (GA₁ and GA₄) were located mainly in theupper part of the hypocotyl and the treatment with uniconazolemarketly reduced the endogenous level of gibberellins thereto less than 11% of the control level. These results suggestthat levels of endogenous gibberellins possibly control theresponse to IAA in these segments. (Received May 12, 1994; Accepted November 15, 1994)     

Effects of Interactions between Low-temperature Treatments, Gibberellin (GA3) and Photoperiod on Flowering and Stem Height of Spring Rape (Brassica napus var. annua)

Exogenous gibberellin A₃(GA₃) reduced the number of leaf nodesat flowering and time to flowering and increased the stem heightat flowering in three genotypes of spring rape (Brassica napusvar.annua L.). The responses to GA₃were similar to those forlong days (LD) and low-temperature treatments, suggesting thatthe effect of photoperiod and the vernalization response areprobably mediated through gibberellins. The response to exogenousGA₃was greatest in non-cold-treated plants in short days (SD)suggesting that endogenous GAs are limiting in these conditions.CCC, an inhibitor of gibberellin biosynthesis, caused a smallincrease in the number of leaf nodes at flowering and time toflowering and a small decrease in the stem height at flowering,but unexpectedly, its effect was hardly influenced by the applicationof exogenous GA₃. Genotypes that showed the clearest responsesto the treatments with regard to the number of leaf nodes atflowering and time to flowering did not show the clearest responseswith regard to the stem height at flowering; the pattern ofresponses of the number of leaf nodes at flowering and timeto flowering was distinct from that of stem height at flowering.This indicates that flower formation and stem elongation areseparable developmental processes which may be controlled bydifferent endogenous gibberellins, different levels of a specificendogenous gibberellin, or different responses to gibberellin.Copyright 1999 Annals of Botany Company Brassica napus var. annua, gibberellin, photoperiod, spring rape, vernalization.     

The regulation of leaf elongation and xyloglucan endotransglycosylase by gibberellin in 'Himalaya' barley (Hordeum vulgare L.)

The potential role of xyloglucan endotransglycosylase (XET)in GA-stimulated cell elongation was investigated during leafexpansion in barley (Hordeum vulgare L.). XET activity in aqueousextracts of leaves was detected in all segments along the elongatingblade of leaf 1 of seedlings, but was at highest levels in basalsegments. Leaf 1 elongation rates of gibberellin (GA)-responsivedwarf mutants were lower than the wild type, and accompaniedby reduced levels of XET activity. Leaf elongation rates ofthe dwarfs increased following treatment with gibberellic acid(GA₃) associated with higher levels of XET activity. The slendermutant, crossed into a dwarfing background, exhibited high ratesof leaf 1 elongation and high levels of XET activity withoutadded GA₃. The elongation of leaf 3 in a GA-responsive dwarfmutant was also studied. Treatment with GA₃ resulted in bladeand sheath lengths being 5-fold and 7-fold (respectively) thelengths of controls, and again there were increases in bladeand sheath XET activities. To investigate the basis for changesin XET activity levels two XET-related cDNA clones were isolated.RNAs detected by the two clones occurred at the highest levelsin basal segments of rapidly elongating leaves, but they haddifferent distribution patterns along the leaf. Overall, thedata indicate that an XET-like activity is detectable in barleyleaves, that the activity level and related. Key words: Gibberellin (GA), leaf elongation, Hordeum vulgare, xyloglucan endotransglycosylase (XET)     

Phase Change in Hedera helix L: II. THE POSSIBLE BOLE OF ROOTS AS A SOURCE OF SHOOT GIBBERELLIN-LIKE SUBSTANCES

When synthesis was estimated by the agar diffusion techniqueboth basal and lateral adventitious roots of Hedera helix L.in the juvenile growth phase were shown to synthesize gibberellin-likesubstances. Seedlings and cuttings from juvenile ivy could begrown in water culture for several weeks. When roots were excisedfrom these plants shoot growth was reduced in comparison withthat of intact plants. The stems, apices, and leaves of derootedseedlings and cuttings contained lower levels of extractablegibberellin-like substances than comparable organs of intactplants. The major zone of gibberellin-like activity in intactplants co-chromatographed with gibberellins A₁ and A₃. Whenthese gibberellins were applied to plants grown in culture theywere found to promote growth of intact but not derooted plants.These findings are discussed in relation to the role of rootfactors and particularly gibberellins in phase change.     

Quantitative Analysis of Endogenous Gibberellins in Normal and Dwarf Cultivars of Rice

Endogenous levels of gibberellins in shoots and ears of twodwarf rice (Oryza sativa L.) cultivars, Tan-ginbozu (dx mutant)and Waito-C (dy mutant), were analyzed and compared with thoseof normal rice cultivar, Nihonbare. The endogenous levels of13-hydroxylated gibberellins in Tan-ginbozu were much lowerthan those in Nihonbare. In Waito-C, the levels of GA₁₉ andGA₂₀ in the shoots were higher but that of GA₁ was lower thanthe levels of these gibberellins in Nihonbare. These resultssupport the hypothesis that the dy gene controls the 3ß-hydroxylationof GA₂₀ to GA₁ while the dx gene controls a much earlier stepin the gibberellin biosynthesis. Our results indicate that GA₁is the active gibberellin that regulates the vegetative growthof rice. The endogenous levels of GA₄ in the ears of the twodwarf cultivars of rice were higher than the level of GA₄ inthe ears of the normal cultivar, Nihonbare suggesting that thebiosynthesis of gibberellin is not blocked in the anthers ofthe dwarf rice. (Received April 27, 1989; Accepted July 12, 1989)     

Interactions Between Nitrogen and Manganese Nutrition of Barley Genotypes Differing in Manganese Efficiency

Ammonium-fed plants may acidify the rhizosphere and thus increaseavailability of Mn in calcareous alkaline soils. The importanceof N nutrition in the differential expression of tolerance toMn deficiency among cereal genotypes is not yet clear. Two factorialexperiments testing effects of the NH₄-N/NO₃-N ratio and Mnfertilization on growth of barley genotypes differing in toleranceto Mn deficiency were conducted in two calcareous alkaline soilsin pots in a controlled environment. In the soil containing80% CaCO₃at pH 8.5, better root and shoot growth and highershoot Mn concentrations were achieved with nitrate supply, especiallyat lower rates of Mn fertilization. The Mn-efficient genotypeWeeah (tolerant of Mn deficiency) achieved better root and shootgrowth than Mn-inefficient Galleon barley (sensitive to Mn deficiency)regardless of experimental treatment. Fertilization with Mndid not influence total N concentration in barley roots andshoots. In the soil containing 5% CaCO₃at pH 7.8, ammonium-fedplants had better root and shoot growth and, at shoot Mn concentrationsabove the critical level, Mn-inefficient Galleon performed betterthan Mn-efficient Weeah barley. It appears that differentialexpression of Mn efficiency among barley genotypes is not associatedwith differences in Mn availability expected to be producedby differential rhizosphere acidification as a response to differentforms of N supply. There is an apparent preference of locallyselected barley genotypes for nitrate nutrition when grown onthe highly calcareous alkaline soils of southern Australia. Ammonium; calcareous soil; Hordeum vulgare ; manganese; nitrate; nitrogen form; nutrient efficiency; rhizosphere     

Variations with Age in Net Assimilation Rate and other Growth Attributes of Sugar-beet, Potato, and Barley in a Controlled Environment: With two Figures in the Text

Sugar-beet, potato, and barley plants were grown in a controlledenvironment, for periods of up to 10 weeks from sowing, witha light intensity of 1,8oo f.c. (4·9 cal./cm.²/hr.) anda temperature of 20° C. during the 18-hour photoperiod and15° C. during the dark period, to test whether net assimilationrate varied with age and differed between the three species. Net assimilation rate of all species based on leaf area (E_A)fell approximately linearly with time. During 5 weeks E_A ofsugar-beet decreased by only about 20 per cent. and E_A of potatodecreased by 50 per cent. E_A of barley remained approximatelyconstant for 4 weeks after sowing and was halved during thesubsequent 4 weeks. The average value of E_A for all times wasgreatest for sugarbeet and least for barley. Net assimilation rates based on leaf weight (E_W) and leaf N(E_N) decreased at about 15 per cent. of the initial value perweek for all species; this was similar to the mean rate of decreaseof E_A of potato and barley, but greater than that of E_A of sugar-beet.Mean values of E_W or E_N for potato and barley were similar andless than for sugar-beet. Relative growth rate (R_W), relative leaf growth-rate (R_A), andleaf-area ratio (F) fell with time at similar rates for allspecies. Average values of R_W decreased and of F increased inthe order sugar-beet, potato, barley. R_A was greatest for potatoand least for barley.     

Changing Kinetic Properties of Barley Alcohol Dehydrogenase during Hypoxic Conditions

The Finnish barley cultivar (Hordeum vulgare L, cv. Hankkija-673)was grown in solution culture for periods of one and two months.The one month old plants had seminal roots, whereas the twomonth old plants grew adventitious roots from the lower nodes.The roots were tested for ADH activity during aeration, during3 d of hypoxic treatment (brought about by passing nitrogenthrough the nutrient solution), and during a 4 d recovery period.The ADH activity, calculated on a protein basis, rose about4-fold during the nitrogen treatment, the greatest increaseoccurring in the adventitious roots. Differences in the kineticproperties of ADH during the hypoxic period were also foundbetween seminal and adventitious roots. The K_m for acetaldehydechanged little in the seminal roots during the hypoxic treatment,but in the adventitious roots it decreased considerably. Thephysiological significance of these changes is discussed. Key words: Hordeum vulgare, alcohol dehydrogenase, hypoxia, enzyme kinetics     

The Interaction of Genetic, Environmental, and Hormonal Factors in Stem Elongation and Floral Development of Cucumber Plants

Hypocotyl length was found to vary between cucumber plants carryingdifferent genes controlling sex expression. Among lines havingonly unisexual flowers (genotype M/M), the homozygous monoeciousplants (st⁺/st⁺) had significantly longer hypocotyls than theirgynoecious counterparts (st/st), heterozygous gynoecious plants(st+/st) being intermediate. Similarly, hypocotyls of plantsof an andromonoecious line (st⁺/st⁺ m/m) were significantlylonger than in their hermaphrodite counterparts (st/st m/m).Differences in intemode length were also significant and inthe same direction. Since stem and particularly hypocotyl elongationin cucumber is known to be very sensitive to applied gibberellin,these findings suggest the existence of differences in the effectivelevels of endogenous gibberellins in the different sex types,higher levels being correlated with stronger male tendency.This conclusion is in good agreement with the known effect ofapplied gibberellin on sex expression (enhancement of the maletendency) in cucumber. Application of gibberellin (GA₄+GA₇) and exposure to ‘summer’conditions (long days and relatively high temperature) inhibitedthe development of pistillate flowers while ‘winter’conditions (short days, lower temperature) had a similar effecton staminate flowers. The effect was in either case specific,that is, limited to flower development. It is concluded thatexogenous and endogenous gibberellins affect not only the initiation,but also the further development, of flowers in the cucumber.     

Nitrogen-13 Studies of Nitrate Fluxes in Barley Roots: II. EFFECT OF PLANT N-STATUS ON THE KINETIC PARAMETERS OF NITRATE INFLUX

Influx of nitrate into the roots of intact barley plants wasfollowed over periods of 1–15 min using nitrogen-13 asa tracer. Based on measurements taken over 15 min from a rangeof external nitrate concentrations (0·2–250 mmolm^–3), the kinetic parameters of influx, I_max and K_m, werecalculated. Compared with plants grown in the presence of nitrate throughout,plants that had been starved of N for 3 d showed a significantlygreater value ofI_max for ¹³N-nitrate influx (by a factor of1·4–1·8), but a similar value of K_m (12–14mmol m^–3). Pre-treating N-starved plants with nitratefor about 5 h further increased the subsequent rate of ¹³N-nitrateinflux, but had little effect in the unstarved controls. Allowingfor this induction of additional nitrate transport, the differencein rates of nitrate influx in control and N-starved plants wassufficient to account for the previously-observed differencein net uptake by the two groups of plants. In barley plants grown without any exposure to nitrate, butwith ammonium as N-source, both I_max and K_m for subsequent ¹³N-nitrateinflux were significantly decreased (by about one-half) comparedwith the corresponding nitrate-grown controls. The importance of changes in the rate of influx in the regulationof net uptake of nitrate is discussed. Key words: Ion transport, nitrate, influx, kinetic parameters, N-deficiency     

A Crucial Role for Gibberellins in Stress Protection of Plants

The hypothesis that there is an intimate relationship betweengibberellin levels and plant stress protection has been testedusing near-isogenic lines of a normal and dwarf barley (Hordeumvulgare L.). Application of paclobutrazol (a triazole), inducedstress protection in the normal line and application of GA₃to the dwarf reversed the inherent stress tolerance. Reversalof the dwarf phenotype by specific gibberellins (GAs) suggeststhat the conversion of GA₂₀ to GA₁ and GA₉ to GA₄ has been compromised.These observations indicate that modulation of specific GAsplays a key role in stress protection. The preferential useof non-chemical technologies including enzyme regulation, phytochromeA overexpression, and breeding for induction of stress tolerancein plants are discussed. (Received November 26, 1998; Accepted March 11, 1999)     

Wild Oat/Barley Interactions: Varietal Differences in Competitiveness in Relation to K+ Supply

Eight cvs of barley (Hordeum vulgare L.) were separately plantedwith Wild Oats (Avena fatua L., genetically pure line CS40)in a sand culture with two external K⁺ concentrations. Substantialdifferences were observed among barley cvs in their abilityto compete with wild oat. The variety Fergus was highly competitiveat both high and low [K⁺]_e, whereas Steptoe was competitiveonly at high [K⁺]_e, and Compana was only weakly competitivewith wild oat. The differences between barley cvs were relatedto their previously reported efficiencies of K⁺ uptake and utilization. Hordeum vulgare L., Avena fatua L., barley, wild oat, competition, K⁺ nutrition, utilization efficiency     

The Influence of Plant Nitrogen Status on NO2 Uptake, NO2 Assimilation and on the Gas Exchange Characteristics of Barley Plants Exposed to Atmospheric NO2

Barley plants were grown in nutrient solution at two contrastingnitrate concentrations to produce plants of low or high nitrogen(N) status. Leaves were then exposed continuously to either0.3 mm³ dm^–3 NO₂ or clean air, with the roots and rootingmedium isolated from the polluted air. Uptake of NO₂ was measuredin two ways; as depletion from an air stream containing thegas and using ¹⁵N-labelled NO₂. Results from the two methodsagreed well and demonstrated that the flux of NO₂ into the leavesof N-deficient barley was lower than that of N-sufficient plants.Nevertheless, the relative contribution of¹⁵N derived from ¹⁵NO₂to the N status of the plant was greater in the plants suppliedwith low nitrate. A major factor in regulating NO₂ uptake bybarley leaves appeared to be stomatal conductance, althoughinternal conductance may also be involved. The effects of NO₂exposure of barley on carbon dioxide exchange rates, transpirationand water vapour conductance were also influenced by the N statusof the plant. Key words: Hordeum vulgare, ¹⁵N-labelled NO₂, carbon dioxide exchange, transpiration     

Gibberellins in Light-Grown Shoots of Pisum sativum L. and the Influence of Reproductive Development

The presence of GA₉, GA₁₉ and GA₂₀ was demonstrated by gas chromatography/massspectrometry (GC/MS) and the presence of GA₄₄ strongly indicatedby GC/MS in selected ion monitoring mode (GC/SIM) in extractsof shoots of light-grown tall peas (Pisum sativum L.). Usingthe rice seedling bioassay with cv. Tan-ginbozu, the levelsof gibberellins in pea shoots were monitored from early shootgrowth through to apical senescence in a tall pea line. Levelsof activity corresponding to GA₂₀, GA₁₉ and GA₄₄ remained relativelystable in the shoot despite reproductive development and apicalsenescence. The level of GA₁-like activity increased to a maximumwhen the leaves had between 7 and 9 leaves expanded and decreasedonly with apical senescence. The na gene which blocks the productionof biologically active gibberellins in shoots but not in developingseed, was also operative in pod walls, with na pods containinglittle or no significant gibberellin-like activity when comparedto na pods at contact. This occurred despite the presence ofrelatively high levels of gibberellins in developing seed atthe same time. The results suggest that there is little or nosignificant leakage of biologically active gibberellins fromdeveloping seed to pods or shoots. Extracts of pods of tallpeas with Na contained low levels of gibberellin-like activitybut like developing seed, contained little or no significantGA₁-like activity despite the presence of significant GA₁-likeactivity in shoot extracts of tall peas. (Received March 11, 1986; Accepted May 27, 1986)     

Mitotic Activities and Levels of Nuclear DNA in the Apical Meristem of Silene armeria (strain SI.2) Following Application of Gibberellin A3

Strain S1.2 of Silene armeria was grown under an 8h-photoperiodand treated with GA₃ every day for 20 days. This growth substancecaused stem elongation, but no flowering in this long-day plant.Changes in the mitotic index and DNA content of cells in thevarious zones of the apical meristem before, during and afterGA₃ treatment were described. Mitotic activity and increasein the proportion of nuclei at the 4C level (S+G₂ phase of thecell cycle) were strongly stimulated in the rib-meristem, andto a lesser extent in the lateral zone, but not in the axialzone. This stimulation of apical activity reached a peak aftertwo GA₃ treatments and then declined gradually, so that after20 days the activity in GA₃-treated meristems was lower thanthat in untreated controls; at this point most cells were inthe G₁ phase. When the GA₃ treatment was discontinued, there was a gradualincrease in the mitotic activity which ultimately reached thesame level as that in controls. Stem elongation ceased and leavesformed aerial rosettes. It is concluded that in vegetative plants of strain S1.2 ofSilene armeria GA₃ acts mainly on the rib-meristem cells whichresults in stem elongation. Lack of response in the axial cellsexplains why GA₃ fails to induce flowering in this strain ofSilene armeria. (Received June 18, 1983; Accepted August 3, 1983)     

The effects of temperature and the Rht3 dwarfing gene on growth, cell extension, and gibberellin content and responsiveness in the wheat leaf

The effects of low temperature and the Rht3 dwarfing gene onthe dynamics of cell extension in leaf 2 of wheat were examinedin relation to gibberellin (GA) content and GA-responsivenessof the extension zone. Leaf 2 of wild-type (rht3) wheat closelyresembled that of the Rht3 dwarf mutant when seedlings weregrown at 10C. The maximum relative elemental growth rate (REGR)within the extension zone in both genotypes was lower at 10Cthan at 20C, but the position with respect to the leaf basewas unaffected by temperature. The size of the extension zoneand epidermal cell lengths were similar in both genotypes at10C. Growth at 20C, instead of 10C, increased the lengthof the extension zone beyond the point of maximum REGR in thewild type, but not in the Rht3 mutant. Increasing temperatureresulted in longer epidermal cells in the wild type. Treatingwild-type plants at 10C with gibberellic acid (GA₃) also increasedthe length of the extension zone, but the Rht3 mutant was GA-non-responsive.However, the concentrations of endogenous GA₁ and GA₃ remainedsimilar across the extension zone of wild-type plants grownat both temperatures, despite large differences in leaf growthrates. The period of accelerating REGR as cells enter the extensionzone, and the maximum REGR attained, are apparently not affectedby GA. It is proposed that GA functions as a stimulus for continuedcell extension by preventing cell maturation in the region beyondmaximum REGR and that low temperature increases the sensitivitythreshold for GA action. Key words: Cell extension, gibberellin, Rht3 dwarfing gene, temperature, wheat leaf     

Growth Hormones and the Creeping Thistle Rust

A re-examination of the life cycle of the rust fungus Puccimapunctiformis (Str.) Rohl. (syn. P. suaveolens (Pers.) Rostr.,P. obtegens Tul. ex Fuckel) on the creeping thistle Cirsiumarvense (L) Scop, has led to the development of a method forgrowing comparable healthy and diseased thistle plants in thegreenhouse. These were used to define the developmental changeswhich result from systemic infection. Closely similar changesare produced by the exogenous application of gibberellic acid(GA₃). The endogenous gibberellin content of systemically diseasedthistle plants is consistently higher than that of healthy plantsduring the earlier part of the season when they are growingfaster. Evidence is given for the presence of two gibberellinswhose relative proportions change during development. The firsthas not been characterized but there is some evidence that thesecond may be GA₃. These results support the hypothesis thatsome of the characteristic features of systemic infection ofthe host are due to the excess production of gibberellins.     

The Relationship of Endogenous Gibberellins to Light-regulated Stem Elongation Rates in Dwarf and Normal Cultivars of Pisum sativum L.

Dwarf cultivar Progress No. 9 and normal cultivar Alaska ofPisum sativum L. were grown under conditions of darkness orred light. Red light decreased the stem elongation rate of bothcultivars. Gibberellins present during the linear phase of stemelongation were isolated and the two main components were tentativelyidentified by gas chromato-chromatography and mass spectrometryas Gibberellin A₁ and A₅. Both gibberellins varied quantitativelywithin and between cultivar and treatment groups, and the amountspresent were inversely related to stem elongation rate. Althoughthe stem elongation rate of plants grown under red light wasrepressed, endogenous gibberellin was not limiting and was asmuch as two-fold higher in red light-grown plants than in dark-grownplants. The levels of endogenous gibberellin in dawrf plantsindicated that the genetic growth limitation was not due toa gibberellin deficiency. (Received May 26, 1981; Accepted January 28, 1982)