Genotypic Differences in Leaf Water Relations between Brassica juncea and B. napus

Various kinds of measurement of tissue water status were madeseveral times during water stress and recovery in Brassica juncea(cv Canadian Black) and B napus (cv Drakkar) Unstressed plantsof the two species had similar leaf water potentials (_w), solute(_s) and turgor potentials (_p) Values of relative water content(RWC) and the slope of the linear relationship between _p andRWC (_p/RWC) were greater in B napus than in B juncea Statistical correlations of pooled data for the watered andstressed treatments differentiated the relationships among RWC,_w and its components in the two species The major statisticaldifference was that _p/RWC was related to RWC in B napus andto _w and _s in B juncea A decline in _p/RWC with decreasing _sin B juncea may be a mechanism for maintaining _p at low soilwater potentials through maintenance of more elastic cell walls. Brassica juncea, Brassica napus, osmotic adjustment, tissue elasticity, water relations     

Photosynthetic Oxygen Evolution at Low Water Potential in Leaf Discs Lacking an Epidermis

Land plants encountering low water potentials (low _w) closetheir stomata, restricting CO₂ entry and potentially photosynthesis.To determine the impact of stomatal closure, photosyntheticO₂ evolution was investigated in leaf discs from sunflower (Helianthusannuus L.) plants after removing the lower epidermis at low_w. Wounding was minimal as evidenced by O₂ evolution nearlyas rapid as that in intact discs. O₂ evolution was maximal in1 % CO₂ in the peeled discs and was markedly inhibited when_w was below –1·1 MPa. CO₂ entered readily at all_w, as demonstrated by varying the CO₂ concentration. Resultswere the same whether the epidermis was removed before or afterlow _w was imposed. Due to the lack of an epidermis and readymovement of CO₂ through the mesophyll, the loss in O₂ evolvingactivity was attributed entirely to photosynthetic metabolism.Intact leaf discs showed a similar loss in activity when measuredat a CO₂ concentration of 5 %, which supported maximum O₂ evolutionat low _w. In 1 % CO₂, however, O₂ evolution at low _w was belowthe maximum, presumably because stomatal closure restrictedCO₂ uptake. The inhibition was larger than in peeled discs at_w between –1 and –1·5 MPa but became thesame as in peeled discs at lower _w. Therefore, as photosynthesisbegan to be inhibited by metabolism at low _w, stomatal closureadded to the inhibition. As _w became more negative, the inhibitionbecame entirely metabolic.     

Water-Relations Parameters of the Phloem. Determinations of Volumetric Elastic Modulus and Membrane Conductivity using an Applied Force Method and Shrinkage and Swelling of Tissues in Solutions at Differing Osmotic Pressure

Water-relations parameters were measured on sections of secondaryphloem from red oak (Quercus borealis michx. f.) and white ash(Fraxinus americana var. biltmoreana [Beadle] J. Wright) usinga linear displacement transducer. Changes in tissue thicknessin response to changes in the osmotic pressure of the bathingsolution were used to calculate the volumetric elastic modulusplus osmotic pressure (_v + ) of the tissue, and an applied forcemethod was used to estimate the time constant for water equilibration(T). The hydraulic conductivity of the cell membranes (L_p) wascalculated utilizing _v + and r values. The time-dependent behaviour of the tissue was much more complexthan originally expected. A correction for a time-dependentprocess that we call ‘drift’ was required to obtainnumbers for _v + . Furthermore, _v + was calculated on two assumptionsin order to relate changes in tissue dimensions to sieve elementparameters. In the first case, a lower limit for _v + of thesieve elements was determined by attributing all changes intissue dimensions to these cells. For red oak the average _v+ on this assumption is 72 bars. Assuming that all cell typeswere equally responsible for the changes in tissue dimensionsresulted in an _v + value of 192 bars for oak. If _v + and rare the same for all cells in the tissue, L_p for the sieve elementsof oak is 9.6 x 10^–8 cm s^–1 bar^–1. Exudationfrom the sieve elements of white ash during excision of thephloem led to artificially high values of _v + for that species. Quercus borealis michx. f., Fraxinus americana var, biltmoreana (Beadle) J. Wright, red oak, white ash, water relations, phloem, volumetric elastic modulus, membrane hydraulic conductivity     

Abscisic Acid and Water Relations of Rice (Oryza sativa L.): Sequential Responses to Water Stress in the Leaf

Water stress was imposed by withholding water at an early vegetativestage from plants of two rice cultivars (IR20 and 63–83)grown in pots. As stress intensified the following sequenceof responses of the leaves was observed: (i) rise in abscisicacid (ABA) content, (ii) closure of stomata, (iii) initiationof leaf rolling. In both cultivars, turgor (_p) declined linearly with total waterpotential () of the leaf. Bulk leaf ABA content increased linearlyas _p declined, and attained twice the control (unstressed) levelfollowing a reduction in _p of about 0.12 MPa. Stomatal conductance exhibited a sigmoidal relationship to _p,declining abruptly when a particular ‘critical’_p was reached (threshold response). The critical potentialsvaried considerably between experiments, but were closely correlatedwith control potentials and with the potentials at which ABAconcentration doubled relative to controls. Leaf rolling was initiated at s near to zero _p. Increases inthe ratio of adaxial to abaxial conductance were associatedwith rolling. Variations in the above responses could be accounted for byvariations in the rate of stress development, which in termsof reduction ranged from 0.38 to 0.86 MPa day^–1. Fastdrying rates resulted in: (a) reduced osmotic adjustment, (b)increased amounts of ABA in the leaf at a given level of or_p, (c) an increase in the ABA concentration present at 50 percent stomatal closure, and (d) initiation of leaf rolling ata higher . Oryza sativa L., rice, water stress, stomata, leaf rolling, abscisic acid     

Osmotic Properties of Drought Stressed Periwinkle (Catharanthus roseus) Genotypes

The pressure-volume technique was employed to compare waterrelations and moisture stress-induced osmotic adjustment ofPeriwinkle (Catharanthus roseus) cv. Pink (PC), Oscillatus (REC)and White (WC). Leaf water potential (_w), osmotic potential(_s), turgor potential (_p), bulk modulus of elasticity (), boundwater (RWC_w) and leaf hydration (H), were estimated by exposingthe plants to a drying cycle during which well watered plantswere dehydrated to zero turgor, and then irrigated. Osmoticadjustment (_w ¹⁰⁰) was calculated by comparing _a at full hydration(_a ¹⁰⁰) in stressed plants after recovery, with _a ¹⁰⁰ in controlplants. Values of ₂¹⁰⁰ were 0.76, 0.33 and 0.11 MPa in cv. PC,REC and WC, respectively. Maintenance of _p at lower ₃ and relativeleaf water content (RWC) in prestressed PC was attributableto a higher alkaloid content and greater leaf cell wall elasticity.RWCW was plotted against _p to determine its contribution tohydration maintenance at lower _p. Genotype PC showed greaterRWC_w at lower _p compared with REC and WC. The present studyhas demonstrated that there are cultivar differences in alkaloidaccumulation and water relations in acclimated plants and thatthe relative ranking for drought resistance within periwinkleappeared to correspond with the changes in osmotic properties. Medicinal plant, drought resistance, alkaloids, periwinkle [Catharanthus roseus (L.) G. Don]     

Wilting of Leaves in Vicia faba L.

Wilting of leaves of Vicia faba L., which occurs when the pressurepotential (_p) is zero, and the leaf-water potential () at wiltingboth depend entirely upon the solute potential at incipientplasmolysis (_so) and not on soil-water status. Wilting in V.faba is acropetal; this is consistent with the hypothesis thatthere is a gradient of decreasing _so up the plant and that wateris transferred from the lower to the upper leaves, hasteningthe overall water loss from the lower leaves to the point when_p is zero. The gradient in _so up the plant is of the order of3–8 bar. It is proposed that wilting when _p>0 (i.e. > _so) shouldbe ‘apparent wilting’ and that when _p0 (i.e. _so),‘true wilting’.     

Alternative Methods of Analysing Water Potential Isotherms: Some Cautions and Clarifications: I. THE IMPACT OF NON-IDEALITY AND OF SOME EXPERIMENTAL ERRORS

In recent years alternative ways have been proposed to transformmeasurements of leaf water potential, , and relative water content,R^*, in order to derive values of osmotic pressure at full turgidityin leaves and shoots, ^o(when 0). Two types of transformationsare usually considered: 1/ versus R^* and versus 1/R^*, and linearregression is used to fit the data in the region where turgoris thought to be zero. It appears that when ^o is estimated bylinear extrapolation of 1/Psi; versus R^* then apoplastic watermight not influence the accuracy of ^o but when the versus \/R^*transformation is used apoplastic water causes an underestimateof ^o. We examine the accuracy of the estimate of ^o obtainedfrom the two transformations when there are random errors in, systematic errors in , and when the osmotic solutions arenon-ideal. The 1/ versus R^* transformation generally producesthe best estimate of ⁰ by linear extrapolation.     

Stomatal Response to Water Stress and its Relationship to Bulk Leaf Water Status and Osmotic Adjustment in Pearl Millet (Pennisetum americanum [L.] Leeke)

The water potential () at which stomata completed closure (_8Lmin)was determined for pearl millet (Pennisetum americanum [L.]Leeke) at two growth stages by monitoring changes in leaf conductance(g_L) and following shoot detachment. Leaf water status wasevaluated concurrently using a pressure-volume (P-V) technique. In a pot experiment with young vegetative plants, _8Lmin closelyapproximated to the estimated at zero turgor (_u) both for controland for drought-conditioned plants which had osmotically adjusted.However, for penultimate leaves of field-grown flowering plants,_8Lmin was found to be 0.61 (irrigated plants) and 0.87 (droughtedplants) MPa below _u. In drought-stressed field-grown plants,osmotic adjustment (characterized by a decrease in solute (osmotic)potential (_s ) at both full hydration and zero turgor) was insufficientto maintain a positive bulk leaf turgor potential (_p) once had declined to below about -1.5 MPa. It is suggested that localizedadjustment by the stomatal complex in response to environmentaldifferences, leaf ageing and/or ontogenetic change, is responsiblefor the uncoupling of stomatal from bulk leaf water status. Key words: Stomata, Water stress, Pennisetum americanum     

Stomatal Behaviour and Water Relations of Chilled Phaseolus vulgaris L. and Pisum sativum L.

Leaf diffusion resistance interpreted as stomatal resistance,leaf water potential (_w), solute potential (_s) and leaf turgorpotential (_p) of the chilling sensitive species Phaseolus vulgariswere determined during chilling at 4 °C in the light. Bothchill-hardened and non-hardened plants were used. For comparison,the chilling resistant species Pisum sativum was also used. The results for chilled P. sativum were similar to those obtainedfor chill-hardened P. vulgaris plants receiving a chilling treatment.In both cases a reduction in stomatal aperture and the maintenanceof a positive leaf turgor were the responses to chilling. Leavesof chilled but non-hardened P. vulgaris plants were found tomaintain open stomata throughout the chilling treatment despitea severe wilt developing after 7 h at 4 °C. This was incontrast to the chill-resistant P. sativum. which showed a rapidclosing and subsequent re-opening of the stomata to a new reducedaperture. During the first 12 h of chilling _wof P. vulgaris leaves changedfrom –0.47 MPa to –1.24 MPa. On more prolonged chilling_w tended to return to pre-chilling values. In addition. _p decreasedfrom 0.42 MPa to zero after only 9 h of chilling, and remainedat this value for the remainder of the chilling period, _s, changedrapidly from –0.89 MPa to –1.35 MPa in the first7.5 h, and after 9 h. _w and _s, were equal, i.e. zero _p. In contrast,the chilling resistant plant P. sativum maintained a positive_p throughout the chilling period, and there was little differencebetween values of _w, and _s in control and chilled leaves. Key words: Chilling, Stomata, ater relations, Phaseolus vulgaris, Pisum sativum     

Water Relations of Sitka Spruce Seedlings After Root Damage

Sitka spruce[Picea sitchensis(Bong.) Carr] seedlings were subjectedto varying degrees of root damage in a growth room, rangingfrom careful transplanting to exposure of the root system toair for up to 3 h. After replanting, transpiration (E), leafwater potential (₁) and growth of the shoot and root were measuredand observations made on plant survival. Some plants in the root exposure treatments died 20–85days after planting. In plants which eventually died, E wasdepressed directly after treatment, but ₁ showed a variableresponse. In some plants ₁ decreased from —8·0x 10⁵ to —30 x 10⁵ Pa after only 10 days but in othersthere was little change in ₁ for 50 days. In spite of the maintenanceof a high water potential in some of the latter plants for longperiods, no root or shoot growth occurred. In plants which lived, the root damage reduced root and shootgrowth relative to untreated controls, and most treatments stronglydepressed E but had little or no effect on ₁. The changes of E and ₁ in treated plants suggest that the suppressionof E was often independent of ₁ although water stress eventuallydeveloped in some of the severely treated plants. Sitka spruce, Picea sitchensis (Bong.)Carr, water relations, root damage, transpiration, leaf water potential     

The Relationship Between Transpiration Rate, Water Potential, and Resistances to Water Movement in Sunflower (Helianthus annuus L.)

The effects of transpiration rate on the vertical gradientsof leaf and stem xylem water potential ( and ) were examinedusing hydroponic sunflower plants. Transpiration was variedby stepwise alterations of environmental conditions. The gradientsof and were relatively small (2.3 and 0.8 x 10⁵ Pa m^–1)when transpiration rates approached zero, but increased sharplyto 5.4 and 2.3 x 10⁵ Pa m^–1 as transpiration increased.However, the gradients were independent of transpiration ratesabove 0.4 g dm^–2 h^–1 owing to variability of theplant resistance. The gradients of _I were usually less thanhalf those of _I. ₁ in individual leaves remained constant over a wide range oftranspiration rates (0.4—2.4 g dm^–2 h^–1) andeach leaf possessed a characteristic plateau value related toits elevation. _I responded similarly but was approximately 2.0x 10⁵ Pa higher than _I at the same elevation. Identical resultswere obtained regardless of the procedure employed to vary transpiration. The drop in water potential between stem and leaf implies thatthe leaf resistance is appreciable. This was confirmed usingrapidly transpiring excised leaves freely supplied with water._I increased by 2.0–2.5 x 10⁵ Pa following removal of theroot resistance but remained 2 x 10⁵ Pa lower than similar excisedleaves in darkness. Furthermore, ^I in excised leaves remainedconstant over a wide range of transporting rates, demonstratingthat the leaf resistance is also variable. The results are discussed in relation to previous reports.     

{gamma}-Amiobutyric Acid Metabolism in Plants: Part 1. Metabolism in Yeasts

The metabolism of -aminobutyric acid (AB) by two yeasts, Saccharomycescerevisiae and Torulopsis utilis, was investigated. Both yeastsgrew well upon AB as a sole source of nitrogen (N), and thelag phase for Torulopsis was shorter than when provided the N-source. The metabolism of AB by Torulopsis, whichwas associated with an increased O₂ uptake, was adaptive incharacter. The enzyme whose formation was induced by the supplyof AB was a transaminase, which was apparently specific forAB as the amino donor. Small amounts of transaminase were presentin unadapted, -grown cells. The optimum pH, equilibrium constant, Michaelis' constant, and coenzyme requirementwere investigated for the transamination reaction involving-ketoglutaric acid (KG) as amino group acceptor. Succinic semi-aldehyde(SSA) was a product of this transamination reaction.The possibility;that some AB was converted into SSA by a direct oxidative deaminationremained unconfirmed. The further conversion of SSA into succinic acid was establishedusing intact. cells for both yeasts. This oxidation processwas shown to be linked to the reduction of pyridine nucleotidesvising extracts of Saccharomyces as a source of SSA dehydrogenase.Dehydrogenase activity could be ascribed to two separate enzymes,one linked to DPN, and the other utilizing TPN and requiringMg⁺⁺ as an activator. The properties of the former enzyme, whichwas more important quantitatively, were investigated and comparedwith those described in the literature for an aldehyde dehydrogenaseof baker's yeast and for SSA dehydro-genases of Pseudomonas.Torulopsis extracts could catalyse the reduction of SSA to -hydroxybutyricacid (OHB); the OHB dehydrogenase involved required TPNH asa coenzyme. Certain other properties of this enzyme are recorded. The possibility is discussed that AB and SSA act as intermediatesin a metabolic pathway that may form a by-pass of the KG-succinatestage of the tricarboxylic acid cycle.     

Natural Abundance of {delta}15N Confirms Insectivorous Habit ofRoridula gorgonias, Despite it Having No Proteolytic Enzymes

Natural abundance values of plant ¹⁵N give an indication asto the source of nitrogen. In particular, carnivorous plantsare expected to be relatively enriched due to trophic enrichmentof their prey. Values of ¹⁵N for adultRoridula gorgonias(mean+3.02)are 4–9 greater than co-occurring non-carnivorous plantspecies and 5.24 greater than juvenileR. gorgoniasplants. Theyare also 3.5–4.26 greater than co-occurringDroseraspecieswhich, being sundews, are considered to be carnivorous. Thesehigh levels of ¹⁵N in adult plants are best explained as beingdue to access to trophically enriched N from insects. As isthe case for other carnivorous plants, leaves and stems ofR.gorgoniasare highly ultraviolet reflective and are thereforeprobably attractive to potential insect prey. This is furthersupport for this plant species being insectivorous.Copyright1998 Annals of Botany Company Nitrogen isotopes, carnivorous plants, insectivorous plant, ultraviolet,Roridula gorgoniasL.     

The Meaning of Matric Potential

The commonly used equation, = P - + , which describes thepartitioning of plant water potential, , into components ofhydrostatic pressure, P, osmotic pressure, , and matric potential,, is misleading. The term , which is supposed to show the influenceof a solid phase on , is zero if a consistent definition ofpressure is used in the standard thermodynamic derivation. However,it can be usefully defined by = + _D, where _D is the osmoticpressure of the equilibrium dialysate of the system. The practicaland theoretical significance of this definition is discussed.     

Water Relations of Potato Leaves. II. Pressure--Volume Analysis and Inferences About the Constancy of the Apoplastic Fraction

Water relations characteristics of potato (Solanum tuberosumL. cv. Bintje) leaves were determined from pressure—volumeanalysis using a pressure chamber. Turgor was 077 MPa and thebulk volumetric modulus of elasticity 81 MPa at full turgidity;turgor loss occurred when water potential () had declined to–087 MPa at a relative water content (RWC) of 0912;the apoplastic water fraction (A) was 0235. As is usually found,there was a linear relation between 1/ and RWC beyond turgorloss. This finding supports the assumptions of the constancyof A during leaf dehydration. Beyond turgor loss the difference between and [measured afterfreezing and thawing (_d)] was about 01 MPa. This differencedid not increase as the leaf water content decreased. This resultcontradicts the constancy of A. It was concluded from calculations with a simple model of leafdehydration that analysis of the relation between and _d providesmore insight in the changes in the apoplastic fraction thanthe relation between 1/ and RWC. Research on the size of theapoplastic fraction and its changes with water potential wouldcomplement current understanding of leaf water relations. Solanum tuberosum, L., water potential, pressure chamber, osmotic potential, pressure potential, relative water content, apoplast, symplast     

Isolation of Subunits of Coupling Factor 1 from Maize and Spinach Chloroplasts and Properties of Combinations of Subunits with ATPase Activity

Subunits (, ß, ) and mixtures of subunits ( ß, , ß , ß ) were isolated without denaturationfrom a chloroform extract of chloroplast coupling factor 1 (CF₁)from maize (Zea mays var. Ushiku 5-4) and from spinach by fastprotein liquid chromatography (FPLC), on an anion-exchange columnof Mono-Q in the presence of n-octylglucoside (OG) and on achromatofocusing column of Mono-P. The ß -subunitcomplex (CF¹ ß ) was the minimum unit required forATPase activity, as was confirmed by the reconstituted complexof ß and subunits. An subunit isolated from maizeinhibited the ATPase activity of CF₁ ß from bothmaize and spinach. CF₁ ß was found to contain anOG-dependent Mg²⁺-ATPase. The ATPase activity of CF₁ ß required divalent cations, such as Mg²⁺ or Mn²⁺, for its expressionin the presence of OG; its optimum pH was 8.0 and it was markedlyinhibited by NaN₃. The enzyme hydrolyzed ATP in prefernece toGTP but not CTP, UTP, ADP, AMP or pNPP. Lineweaver-Burk plotsof its activity were curvilinear in the range of 0.6–0.7mM ATP.Mg²⁺. ¹Present address: Department of Biology, School of Education,Waseda University, Shinjuku-ku, Tokyo, 160 Japan. (Received February 15, 1989; Accepted April 20, 1989)     

The Incorporation of C14-labelled Substrates into the Amino-acids of Groundnut Plants (Arachis hypogaea)

The amino-acid metabolism of groundnut plants has been studiedwith special reference to -methyleneglutamic acid (-MGA) andy-methyleneglutamine (-MG), constituents not found in the greatmajority of plant species. The sequence in which C¹⁴ from radioactive C¹⁴-carbon dioxideenters the amino-acids of leaves was determined. The patternof labelling was very similar to that found for leaves of otherspecies. The metabolic relationships existing between photosynthesisand amino-acid synthesis therefore do not seem to be affectedby the large quantities of -MGA and -MG present in the leaves.-MGA and -MG only gained traces of radioactivity. Experiments designed to study the incorporation of C¹⁴ fromuniformly labelled C¹⁴-alanine into the amino-acids of roots,immature leaves and cotyledons of seedlings and young plantsindicated that the main site of synthesis of -MGA was the cotyledons. Various C^I4-labelled substrates were fed to germinating seedsand, after a period of growth, the specific activities of theamino-acids of seedlings receiving different treatments weredetermined. Comparison of the specific activities enabled certaindeductions to be made concerning the probable biosynthetic pathwaysleading to -MGA and -MG. The results were consistent with theintact incorporation of pyruvate molecules, or another related3-carbon-atom containing compound, into -MGA.     

A Transient Stimulation of Net Photosynthesis of Rye Leaves by {alpha}-Hydroxy-2-pyridinemethanesulphonic Acid ({alpha}-HPMS) due to Inhibition of Photorespiratory CO2 Release

The effects of -hydroxy-2-pyridinemethanesulphonic acid (-HPMS)upon net photosynthesis (P_n, the CO₂ compensation point (),post-lower illumination burst of CO₂ (PLIB) and post-lower temperatureburst of CO₂ (PLTB) in detached rye (Secale cereale L.) leaveswere investigated. At low concentrations ( 0.5 mol m^–3),-HPMS initially stimulated P_n and decreased the magnitude ofboth PLIB and PLTB. The decreased at all concentrations of-HPMS (0.05–5.0 mol m^–3. The effects of -HPMS onP_n and were time-dependent and, after a few minutes, the P_nwas inhibited while values increased considerably. At a higherconcentration (5.0 mol m ^–3), the transient effects of-HPMS were shorter () or not observed at all (P_n. Both PLIBand PLTB, when expressed in relation to P_n, increased at higherlevels of this compound. Similar data with respect to the effectsof -HPMS on PLIB and PLTB were found for leaves of dandelion(Taraxacum officinale L.). The results suggest that -HPMS may stimulate P_n by inhibitingphotorespiration, as originally suggested by Zelitch (1966),but only at low concentrations and over a short time span. Thedecrease of PLIB and PLTB values at low -HPMS levels is consistentwith these processes being a residual activity of the glycolatepathway. Key words: CO₂ compensation point, -hydroxy-2-pyridinemethanesulphonic acid, photorespiration, photosynthesis     

Nitrogenous Compounds and Nitrogen Metabolism in the Liliaceae: II. The Nitrogenous Compounds of Leaves of the Genus Tulipa: Environmental Effects on the Composition of Tulipa gesneriana

Because of the predominance of -methyleneglutamine in Tulipagesneriana a quantitative, chromatographic investigation hasbeen made to show the factors which determine the compositionof the alcohol-soluble nitrogen fraction in Tulipa A comparative study of the amino-acids that occur free in theleaves of fourteen different species of Tulipa has been made,-methyleneglutamine varied from as much as 60–70 per cent,of the amino-acid nitrogen down to levels below those whichpermit its chromatographic detection. The effects of temperature during growth and of diurnal variationon the nitrogenous compounds of the leaves of T. getnerianaare described. Excised shoots of T. gesneriana were exposed to light and darknessto affect the balance of protein synthesis and breakdown sothat the concomitant effects on the -methyleneglutamine acidcompounds and other main components of the soluble nitrogenfraction could be determined. Lastly, the relative ease of entry of C¹⁴ from C¹⁴O₂ into thesugars and certain other compounds of T. gesneriana is contrastedwith the relative difficulty with which it enters the -methyleneglutamineof the mature leaves on excised shoots. The occurrence and metabolism of -methyleneglutamine in T. gesnerianaleaves may, therefore, be regarded as additional to and superimposedupon an otherwise typical pattern of nitrogen metabolism: inthese leaves -methylene-glutamine does not assume the role usuallydischarged by glutamine.