Growth-Regulating Substances and the Growth of Tiller Buds in Barley; Effects of Cytokinins

An hypothesis was set up from which it was predicted that applicationof cytokinin to barley seedlings grown without mineral nutrientswould lead to rapid growth of the coleoptile and first leaftiller buds. Application of cytokinins to the leaves was ineffective,but supplying a number of known cytokinins by steeping the rootsof 4 d old seedlings in solution for 4 h led to significantgrowth of the coleoptile bud. Adenine and cytokinin analogueshad no effect. Supplying cytokinins through the roots also furtherenhanced the growth of buds of plants given mineral nutrients.Cytokinin treatment reduced root dry matter, with small reductionsin mean axis length and number of lateral roots. For plantsnot given mineral nutrients reduction in root weight was compensatedby an increase in weight of the aerial parts; however, for plantssupplied with mineral nutrients this was not so and the lowerroot weight resulted in a smaller total plant dry weight. An interpretation of tiller bud growth in terms of control byinteracting effects of mineral nutrition, assimilate supply,and cytokinin availability is proposed.     

Growth-Regulating Substances and the Growth of Tiller Buds in Barley; Effects of IAA and GA3

Application of cytokinin to barley seedlings grown without mineralnutrients leads to rapid growth of coleoptile and first leaftiller buds. IAA and GA₃ cannot substitute for cytokinin inthis effect and applications of TIBA and CCC are also ineffectiveon bud growth. However, when bud growth was promoted, eitherby application of cytokinins or by supplying plants with mineralnutrients, IAA and GA₃ applications caused enhanced tiller growthindicating that these compounds can stimulate growth of activelygrowing buds. The results are compatible with an interpretationwhich stresses the importance of cytokinin availability in determiningtiller bud growth.     

Temperature Controls a Dependence of Barley Plant Growth on Mineral Nutrition Level

The effect of temperature regime on growth and other morphological characteristics of barley plants (Hordeum distichum L., cv. Andrei) as dependent on the level of mineral nutrition was investigated in a controlled experiment. Plants were raised hydroponically at a high (0.22 g/(g day)) and low (0.05 g/(g day)) relative rates of the addition of mineral nutrients (R _A). Mineral nutrients were daily added to the nutrient solutions in exponentially increased amounts to provide steady-state plant growth. At the optimum temperature regime (21/17°C, day/night), the plant relative growth rate (RGR) was proportional to the preset R _A during the entire exponential period. Low R _A led to a decrease in the nitrogen content in plants, plant weight, and respiratory activity, as well as to the increase in the relative root weight. Biomass accumulation at lowered temperature regime (13/8°C) and a high R _A was 1.8-fold lower than at optimum temperature regime. Although under these conditions, the ratio of respiration to gross photosynthesis reduced threefold due to the decrease in the respiration rate, RGR of plants was equal to 0.11 ± 0.02 g/(g day), which was twice lower than the preset R _A. These pointed to the decrease in plant ability to maintain a certain ratio of photosynthesis to respiration within a day. At a deficiency of mineral nutrition and low temperature, RGR reached the preset R _A. Plants adapted to lowered temperature by a shift of the temperature optimum of their metabolism (heat production) to lower values. As a whole, a low variability of such growth parameters as RGR, C/N, and root to shoot weight ratio at different R _A and lowered temperatures testified to the lessening of growth limitation by the mineral nutrition.__________Translated from Fiziologiya Rastenii, Vol. 52, No. 3, 2005, pp. 384–391.Original Russian Text Copyright © 2005 by Garmash.     

Effects of Plant Growth Regulators and Nutrient Supply on Tiller Bud Outgrowth in Barley (Hordeum distichum L.)

Seedlings of spring barley were raised in 100 and 20% nutrientsolution and treated with a foliar application of Terpal, Cerone,TIBA, GA₃ or BAP. The growth of individual tiller buds and tillers,the main shoot and the root system was recorded over the following15 d. Terpal and Cerone stimulated tiller bud elongation within5 d at both nutrient levels and after 15 d the number of emergedtillers was increased at the higher nutrient level. Terpal characteristicallypromoted the growth of secondary tiller buds whereas Ceronepromoted the emergence of the coleoptile tiller; both thesePGRs also retarded the development of the main shoot. TIBA increasedthe number of elongating tiller buds but this did not resultin greater tillering. GA₃ reduced the number of elongating tillerbuds and restricted their growth, especially in the high nutrientregime; this was accompanied by an increase in main shoot elongation.The growth and development of tiller buds was reduced by BAPand the number of emerged tillers was reduced at 15 d in bothnutrient levels; main shoot dry weight and root elongation werealso reduced. The results are considered in relation to theoverall influence of hormonal factors and mineral supply ontiller bud outgrowth. Hordeum distichum, spring barley, tiller bud outgrowth, plant growth regulators, Terpal, Cerone, GA₃, BAP, nutrient supply, apical dominance, TIBA     

A Comparison of Main-stem and Tiller Growth in Barley; Apical Development and Leaf-unfolding Rates

Information is presented on the hierarchical relationship betweenthe stems of Proctor barley grown under controlled conditions.Rates of production of foliar and floral primordia and ratesof leaf unfolding were found to follow the order main-stem >primary tillers > secondary tillers. Apical dome size wasinitially substantially greater for the main-stem than for thetillers due principally to the slower transition of the latterto the reproductive condition; by the time that double ridgesare formed on the tiller apices apical dome sizes are similarto that of the main-stem. Transition to the reproductive conditionoccurs at about the same time for the primary tillers. Thisnear-synchrony results in a smaller number of leaves borne onsuccessive tillers, since these have had less time to initiatefoliar primordia, and a measure of synchrony in awn appearance,since rates of car development are similar for these stems. The nature of the hierarchical relationship between main-stemand tillers is discussed.     

Studies in the Nutrition of Apple Rootstocks: III. Effects of Level of Magnesium Nutrition on Growth, Form and Mineral Composition

One-year rooted shoots of apple M. VII rootstock were grownfor a single season by spraying their roots continuously withnutrient solutions containing <3, 10, 45, and 135 ppm Mg.These treatments respectively produced plants with a severeMg deficiency (Mg₍₀₎), a mild deficiency (Mg₍₁₎), normal foliageand growth (Mg₍₂₎), and a high level of Mg (Mg₍₃₎). The effectof Mg nutrition on the leaves can explain most of the observedeffects on vegetative growth. Depression of growth of Mg₍₁₎)plants was not detected until late in the season although necrosis,typical of Mg deficiency, occurred on the lower leaves by mid-July.This loss of photosynthetic area of leaf was to a large extentcompensated for by increased efficiency of the remaining greenleaves; as the deficiency symptoms progressed, the loss of leaf,and therefore of carbohydrate production, predominated and totalgrowth was affected. The form of the plant was little altered. Symptoms were apparent on leaves of severely deficient Mg₍₀₎plants by late June, and reductions in growth and modificationsin form by August. The poor root systems and thin shoots arelargely attributed to loss of leaf rather than to the low Mgconcentration in these organs. No symptoms of toxicity developed on the Mg₍₃₎ plants althoughby the end of the season the total dry weights were less thanthose of the Mg₍₂₎ controls, and the plants also produced thickershoots. Plant dry weight, and more especially Mg uptake, increasedfrom Mg₍₀₎ to Mg₍₂₎ but above this level the gradient of Mguptake in relation to supply was less steep. The excess Mg takenup at the Mg(j) level led to the relatively lower weights ofthese plants. Much of the excess Mg accumulated in the leaves,especially in the older ones. Mg moved from, or accumulatedin, the lower leaves of the shoot according to the level ofMg supplied to the plants. Thus the shoot apex tended towardsnormality both in the concentration of Mg and in its growth. In general, growth of the plant, and of individual componentparts, governed the uptake of elements other than Mg, althoughmore K, Ca, and Mn and less Fe per unit of plant dry weightaccumulated at the lower levels of Mg than at the higher levels.     

Mineral Nutrition and Plant Growth Response to Climate Change

The limiting factor concept has often been used to describeplant growth responses to altered availability of resources.However, even preliminary experiments, where atmospheric CO₂concentrations and solution mineral concentrations were varied,demonstrated that a more complex concept was required to interpretthe potential effects of climate change and mineral availabilityon plant growth. It is proposed that these resources for plantgrowth may be better viewed as simultaneously limiting. Further,in considering the limitation in plant growth to mineral nutritionit is important to consider both the solution concentrationand the total amount of the individual minerals available tothe plant. Sustaining a positive response to increased CO₂ concentration,for example, requires an increase in plant uptake of the totalamount of minerals. Consequently, it is very difficult to predictthe plant growth response to climate change because of the largeuncertainty about mineral availability. On the one hand, increasedCO₂ concentrations should stimulate nitrogen fixation by bothfree-living organisms and symbiotic systems, and improve soilproperties for mineral availability as a result of increasedorganic matter deposition in the soil. On the other hand, increasedtemperature and altered rainfall patterns may result in increasedlosses of soil minerals. Even the direction in the net changein available soil minerals is unclear. Realistic evaluationsof the effects of climate change on plant growth will be challengedto contend with the large uncertainty and complexities in understandingmineral availability and plant mineral nutrition.     

Effects of Shading the First Leaf of Barley Plants on Growth and Carbon Nutrition of the Stem Apex

Effects of shading the first leaf on development of the apicalregion were investigated by examining the growth of leaf primordiaand the apical dome in the early seedling stages. Shade treatment affects the size of the dome; it was shown thatvalues for height, width, and volume of the dome of 12-day controlplants were always higher than for shaded plants. Primordialgrowth, in terms of length and dry weight, was reduced by shadeand growth in dry weight of the second, third, and fourth leaveswas shown to be dependent on photosynthetic production by thefirst leaf. Incorporation of ¹⁴C in the apical region was detected by autoradiographyon day 6 and increased with age. Transfer of assimilated carbonfrom the first leaf to the apex occurred during the first 3h after exposure to ¹⁴CO₂. On a unit dry-weight basis it isshown that the third and fourth leaves and apex incorporatedproportionately more labelled carbon than the larger older organssuch as the, second leaf. Shade treatment reduced incorporationinto the apical region and this is associated with the failureof the apex to grow over the period up to day 15. Evidence isprovided to show that in control plants the second leaf suppliescarbon to the apex from about day 12. The crucial importanceof the contribution of the first leaf to plant development isdiscussed.     

The Effects of Photoperiod and Mineral Nutrient Supply on Growth and Primordia Production at the Stem Apex of Barley Seedlings

Growth and development of the shoot apex in seedlings of threebarley cultivars was examined in two daylengths (8, 16 h) andat two mineral nutrient levels (x 1, x 0.1). Production of primordiawas greater at the higher nutrient level and in the longer days.The rate of production varied with cultivar but in all casesthe plastochron shortened with transition to spike formation.Early flowering (cv. Clipper) was associated with a high rateof primordial production and early transition to spike formation,late flowering (cv. Proctor) with a low rate of production anda longer vegetative phase. The cultivar Akka showed intermediatecharacteristics. The volume of the apical dome increased linearlywith increasing numbers of primordia, the rate of increase varyingwith cultivar and treatment. Enlargement of the dome was duemainly to increase in cell number. The transition of the apexto produce spikelet primordia occurred with widely differingvolumes of the apical dome, thus invalidating the hypothesisthat transition is dependent upon attainment of a critical domesize. Although both the rate of production of primordia andenlargement of the dome were markedly affected by photoperiod,both were unaffected when the photoperiodic treatment was givendirectly to the shoot apex. It is considered that the fate of a primordium once initiatedis determined by competition for available metabolites betweenit, other primordia and the apical dome. Hordeum vulgare L, barley, apical dome, primordia, plastochron, cell division     

Balance between Potassium and Phosphorus in the Nutrition of Barley: II. Observations on Growth and Uptake

In K x P interaction experiments with barley, treatments canusually be found in which the plants appear to be balanced betweenthe two types of deficiency symptoms. These treatments favourdry weight and grain production. It is not known whether internalparameters of the health of barley tend to have similar optima.From the results of a water-culture experiment, a comparisonwas made of the patterns of response exhibited by various parametersof the health and growth of barley. Since the patterns did notcorrespond with each other very closely, it was concluded thatexternal indications of the balanced state are not necessarilythe product of optimal internal conditions. It was also considered whether a simple definition of a balancednutrient solution could be achieved by reference to relationshipsbetween dry weight and either nutrient supply or nutrient uptake.Such a definition might have helped to reduce the subjectivityin the use of this term, but no basis was found for sound generalizationsalong these lines.     

The Effects of Shade Treatment and Light Intensity on Rihulose-1,5-Diphosphate Carboxylase Activity and Fraction I Protein Level in the First Leaf of Barley

It has been confirmed that shading leaves from day 5 onwardslowers the rate of CO₂ fixation when they are placed in saturatingirradiances. The reduction due to shade treatment is about 46per cent and a similar reduction in maximum chlorophyll contentof the leaf follows shading. Maximum amounts of total solubleprotein and of Fraction I protein are less in shaded leavesthan in control leaves and prolonged treatment leads to a declinein leaf protein content. The relative amounts of different proteinare also affected by treatment; in control leaves Fraction Iprotein accounts for about 45 per cent of the total but in shadedleaves the value is about 30 per cent. Increases and decreasesin leaf protein amount, with concomitant changes in the ratioof Fraction I to total protein can be brought about by removingshades and re-applying them. Such changes can be induced evenin fully expanded leaves in which net protein synthesis is notusually found. Maximal amounts of leaf protein are found in irradiances of60 W m^–2 or more, with lower values at lower light intensities.Where the first leaf is held in a stream of CO₂-free air a lowerlevel of protein is found. This, and the ratio of Fraction Ito total protein, are similar to values for shaded leaves, andsuggest the involvement of photosynthetic carbon fixation indetermining leaf protein amount. A 1:1 linear correlation between amount of Fraction I proteinand RuDP carboxylase activity is shown but the rate of CO₂ incorporationby leaf extracts is 2–3 times greater than that of theintact leaf. The significance of this and the effect of irradianceon leaf protein amount are discussed.     

Immunofluorescent Localization of Plasma Membrane H-ATPase in Barley Roots and Effects of K Nutrition

The plasma membrane H⁺-ATPase (PM-H⁺-ATPase) of barley (Hordeum vulgare L. cv Klondike) roots was assayed by cross-reaction on western blots and cryosections with an antibody against the PM-H⁺-ATPase from corn roots. Under conditions of reduced K availability, which have previously been shown to increase K influx by greater than 25-fold, there were only minor changes detected in PM-H⁺-ATPase levels. Antibody labeling of cryosections showed the relative distribution of PM-H⁺-ATPase among cell types in root tips and mature roots. Epidermal cells, both protoderm and mature root epidermis, including root hairs, had high levels of antibody binding. In mature roots, the stelar tissue showing the highest antibody binding was the companion cells of the phloem, followed by pericycle, xylem parenchyma, and endodermis.     

The Effect of Mineral Nutrition on the Distribution of Growth in Lolium perenne

Ramets of Lolium perenne from the same genotypes were grownin nutrient solutions of two concentrations. Additional mineralnutrition increased the rate of production of new tillers, therate of increase in size of tillers and the rate of productionof new roots, but not the rate of growth of individual mainroots. The genotypes differed considerably from each other intheir rates of growth, especially of shoot growth. When thegrowth-rate of the same population of genotypes was alteredby changing the environmental conditions it was found that thelower the mean rate of shoot growth, the greater its coefficientof variation. Variation in the rate of shoot growth, due tomineral nutrition and genetic constitution of the plants growingin a dilute nutrient solution, was accompanied by a similarbut smaller variation in the number of roots.     

激素对水稻分蘖芽生长和分蘖相关基因表达的调控效应

以水稻品种‘南粳44＇为材料,研究了植物激素对水稻分蘖芽生长及OsTB1、OsD3、OsD10和OsD27等基因表达的调控。结果表明,水稻分蘖芽生长的调控至少存在两条途径,一条通过调控分蘖芽中CTK含量进而调控OsTB1等基因表达来实现,另一条通过调控OsD3、OsD10和OsD27等系列基因表达来实现。外源IAA和GA_3对水稻分蘖芽生长的调控是通过这两条途径共同实现的。     

Physiological Studies in Plant Nutrition: XVI. The Mineral Nutrition of Bracken

Part I of this paper describes culture methods for the productionof large numbers of bracken sporelings of uniform size. Observationson the growth of the bracken plant and results of experimentson the mineral nutrition of the prothallial and sporophyte generationsare presented. The presentation of further growth data and afull analysis of the results is deferred to the second partof this paper.     

Responses of Leaf and Tiller Emergence and Primordium Initiation in Wheat and Barley to Interchanged Photoperiod

The influence of constant (9, 13 and 19 h) and reciprocally-interchangedphotoperiods [at terminal spikelet (TS) or triple mound (TM)]on leaf, tiller and primordium development were examined usingphotoperiod-responsive cultivars of spring wheat, ‘UQ189’ and spring barley, ‘Arapiles’. In bothspecies, constant longer photoperiod reduced the duration fromsowing (S) to double ridge (DR), as expected. However, photoperiodsensitivity was not restricted to this mainly vegetative phase.There was also a marked increase or reduction in the durationof reproductive phases between TS/TM and heading (H) when plantswere transferred to shorter or longer photoperiods respectively,compared with controls. These responses were largely independentof the photoperiod during previous phases although minor effectsof the previous photoperiod were observed. For both species,the time course of leaf emergence was linear, or bi-linear,depending on the final leaf number on the main stem. The rateof leaf emergence was faster for the first six to eight leavesthan for the leaves appearing subsequently. The rate of emergenceof early-formed leaves was independent of photoperiod whereasthe rate of emergence of later leaves varied with photoperiod.Photoperiod also affected the dynamics of tillering. The rateof leaf primordium initiation was little affected by variationin photoperiod, but the rate of spikelet initiation increasedwith increases in photoperiod. The rates of leaf and spikeletprimordium initiation were both substantially higher in barleythan in wheat. The fact that the reproductive phase from TS/TMto H was largely independent of the duration of the previousphase provides evidence that this phase might be geneticallymanipulated to increase the time for floret development andhence grain number. Copyright 2000 Annals of Botany Company Photoperiod, rate of development and leaf emergence, tillering, wheat (Triticum aestivum L.) and barley (Hordeum vulgare L.)     

Effects of Tiller Separation and Root Pruning on the Growth of Lolium perenne L.

Experiments have been carried out to assess the role of inter-tillerinteractions within plants of Lolium perenne L. cv. S24 as factorswhich could play a part in determining the growth-rate of thewhole plant. In order to do this, the effect of tiller separationon plant growth was studied as well as the influence of theremoval of different parts of the root and shoot system. Wheneverplants were supplied with nutrients by means of culture solutions,separation of tillers brought about a marked increase in theamount of dry matter produced. Root pruning and in particularthe removal of root apices brought about similar increases ingrowth. Separation of tillers in soil-grown plants did not bringabout increased growth. These results are interpreted as indicatingthat separation of tillers or the initial removal of root apicesin plants supplied with nutrients by means of culture solution,promotes root branching which brings about an increased netassimilation rate arising from an increase supply of a growth-promotingsubstance, probably a cytokinin, from the root apices. The resultsare not taken as indicative of an intertiller interaction whichrestricts the growth of intact plants.     

The Effects of Mineral Nutrition and Density on Root Interactions in Three Grass Species

Studies were made of competition between Dactylis glornerataL., Holcus lanatus L. and Anthoxanthum odoratum L. The competitiveability of the species depended on the density of planting.D. glomerata was more competitive when planted at high density,whereas H. lanatus was more competitive at low density. A. odoratumwas most competitive at the intermediate density.     

The Effects of NaCl Treatment on Water Relations, Growth, and ABA Content in Barley Cultivars Differing in Drought Tolerance

Changes in transpiration and stomatal conductance and other characteristics of water relations, growth rate, and ABA content have been followed in short- and long-term experiments in two barley cultivars (cv. Michaelovsky and cv. Prairie) with contrasting drought resistance characteristics. The aim of this work was to reveal the importance of stomatal behavior in salt tolerance and also the involvement of ABA in its control. Salinity stress brought about a reduction in stomatal conductance in both cultivars, but the effect was initially more pronounced in the drought-tolerant cv. Prairie than in the drought-sensitive cv. Michaelovsky. The difference between the two cultivars changed with time, and later on transpiration and stomatal conductance became higher in Prairie than in Michaelovsky. In both the short and the long term, the extent of stomatal closure due to salinity correlated with the level of ABA accumulation in the leaves of the plants. Fast stomatal closure was likely to be responsible for growth resumption after an initial arrest by salt treatment and for the maintenance of extension growth later on, thus enabling its higher rate in Prairie than in Michaelovsky plants. Leaves of Prairie accumulated less toxic chloride ions, which may be the result of a lower transpiration rate observed during the first phase of salt treatment. A subsequent increase in stomatal conductance observed in Prairie is likely to ameliorate their gas exchange and maintain photosynthesis and growth. Thus, differences between the cultivars in the stomatal response to salinity changed with time, which may be why there are discrepancies in the attempts to relate stomatal conductance to salt tolerance observed in literature.     

Physiological Studies in Plant Nutrition: XVII. A General Survey of the Free Amino-acids of Barley Leaves as affected by Mineral Nutrition, with Special Reference to Potassium Supply

A survey by means of paper chromatography of the free amino-acidsin barley leaves at different stages of development is described.The plants were grown in sand culture, using a variety of nutrientsolutions, the most important variable being potassium supply.In general, the amino-acid distribution is not greatly affectedby nutritional differences. Potassium deficiency results in much increased contents of theamides, especially asparagine, and reduced amounts of the correspondingacids. Other acids increased under these conditions are serine,leucine, tyrosine, and phenylalanine, but the most characteristicchanges occur in -the basic constituents; lysine and arginineare increased, and an unknown basic substance accumulates verycharacteristically. Accumulation of this substance (probablyan amine) is checked by the presence of either sodium or rubidiumin the culture solution, almost as effectively as by increasedpotassium supply. Extreme potassium deficiency yields chromatogramsclosely resembling those from more moderately deficient plants. When a low rubidium supply accompanies potassium shortage thedisturbances occasioned by potassium deficiency, both in growthand in amino-acid distribution, are diminished, but a higherrubidium supply leads to toxic symptoms in the plant; largeincreases in the contents of the two amides and of their parentacids are then found, this being the only condition encounteredthat leads to simultaneous accumulation of both these typesof substances.