The Effects of Plant Density on Shoot and Leaf Lamina Angles in Lolium multiflorum and Paspalum dilatatum

The effects of plant density on shoot and leaf lamina zenithangles (i e the angles with respect to the vertical) were investigatedin Lolium multiflorum and Paspalum dilatatum plants grown inpots either outdoors or in a glasshouse Tillers appeared abovethe ligule of their subtending leaf with a small zenith anglebut, in plants grown at low densities, gradually adopted a morehorizontal position In plants grown at high densities this gradualincrease in shoot zenith angle with age was strongly reducedThe average shoot zenith angle at a given time was lower (ie tillers were more erect) at high, compared to low plant densitiesShoot number per plant and average shoot zenith angle per plantwere directly related In P dilatatum plants the leaf lamina remained nearly horizontalat any plant density This observation indicates that the movementof the leaf lamina, pivoting at the point of attachment to theleaf sheath, compensated the movement of the shoot, pivotingat the base of the leaf sheath-tube Shoot angle responses to plant density would place leaf laminaeat different heights within the canopy, affecting their abilityto compete for light and their susceptibility to wind impactand herbivore grazing Foliage architecture, Lolium multiflorum Lam, Paspalum dilatatum Poir, plant density, neighbour effects, shoot angle, leaf angle     

Effects of Chlorocholine Chloride and Water Regime on Growth, Yield, and Water Use of Spring Wheat

When CCC was applied as a spray to the leaves of Brassica oleraceaL. (Brussels sprout) grown in pots, plant height and mean internodelength were reduced. The effects appeared more slowly and wereless pronounced than those previously observed when CCC hadbeen applied to the soil; other differences were that root growthwas not inhibited, stem weight was only significantly reducedat the highest rate of application (2 per cent), and stomatalnumber per unit area of lower leaf epidermis was not affected.In common with soil applications, leaf thickness, stem diameter,and the percentage moisture contents of the leaves were allincreased by foliar applications.In a further experiment theprogress of wilting was observed in untreated plants and inplants treated with CCC applied either to the leaves or to thesoil. The rates of water loss and the moisture contents of theleaf laminae of the treated plants, after a period of wilting,were not significantly different from the controls. The treatedplants, however, looked less ‘wilted‘ for the changein angle of the leaf lamina to the stem was less and their leaveswere therefore held more upright.     

The Effects of Foliar Applications of (2-Chloroethyl)-trimethylammonium Chloride on Leaf Area and Dry Matter Production by the Brussels Sprout Plant

When CCC was applied as a spray to the leaves of Brassica oleraceaL. (Brussels sprout) grown in pots, plant height and mean internodelength were reduced. The effects appeared more slowly and wereless pronounced than those previously observed when CCC hadbeen applied to the soil; other differences were that root growthwas not inhibited, stem weight was only significantly reducedat the highest rate of application (2 per cent), and stomatalnumber per unit area of lower leaf epidermis was not affected.In common with soil applications, leaf thickness, stem diameter,and the percentage moisture contents of the leaves were allincreased by foliar applications.In a further experiment theprogress of wilting was observed in untreated plants and inplants treated with CCC applied either to the leaves or to thesoil. The rates of water loss and the moisture contents of theleaf laminae of the treated plants, after a period of wilting,were not significantly different from the controls. The treatedplants, however, looked less ‘wilted‘ for the changein angle of the leaf lamina to the stem was less and their leaveswere therefore held more upright.     

Salt Tolerance in the Halophyte Suaeda maritima L. Dum.: Intracellular Compartmentation of Ions

The time-course of exchange of sodium and potassium ions fromroot and leaf material of the halophyte Suaeda maritima hasbeen followed and the data analysed according to the phenomenologyof efflux, or compartmental, analysis. Sodium ions were exchangedmuch more slowly (c. 4 times) from the vacuoles of leaf cellsof plants grown in sodium chloride than were potassium ionsfrom the vacuoles of leaf cells of plants grown either in similarconcentrations of potassium chloride or in low concentrationsof potassium. In plants grown in sodium chloride, sodium ionswere exchanged 9 times more slowly from the vacuoles of leafcells than from the vacuoles of root cells. The concentration of sodium ions in the cytoplasm of leaf cellsof plants growing in 340 mol m^–3 sodium chloride was estimatedto be 165 mol m^–3 when the average concentration in theleaf tissue was about 600 mol m^–3. As measured by movement from mature to developing leaves inintact plants; there was less in vivo retranslocation of ²²Naand ³⁶CI in plants growing in sodium chloride than there wasof ⁸⁶Rb in plants growing either in potassium chloride or innon-saline conditions. The results are discussed in terms of the concept and energeticsof compartmentation of ions in the cells of halophytes.     

The Effect of Light Intensity during Growth on the Subsequent Rate of Photosynthesis of Leaves of Tall Fescue (Festuca arundinacea Schreb.)

Leaves of tall fescue (Festuca arundinacea Schreb.) plants grownin bright light had higher rates of apparent photosynthesisper unit leaf area in bright light, and slightly lower ratesin dim light than did those of plants grown in dim light. Darkrespiration rates were higher in plants grown in bright lightthan in plants grown in dim light and the decline of photosynthesiswith increasing leaf age was faster. The rate of apparent photosynthesis in bright light of the firstleaf to become fully expanded after plants were transferredfrom bright to dim light was lower than that of plants remainingin bright light. The decline in the rate of photosynthesis ofa leaf already fully expanded at the time of transfer was notaffected. Transferring from dim to bright light increased therate of photosynthesis of the next expanded leaf; it also increasedthe rate of an already fully expanded leaf during the firstweek in bright light. After this, photosynthesis fell at a ratesimilar to that of plants remaining in dim light.     

Starch Synthesis from Exogenous Sugars in Tobacco Leaf Discs

Sets of discs were taken from leaves of destarched tobacco plants(Nicotiana tabacum L. cv. xanthii) and floated on solutionsof sucrose or glucose in the dark. Abundant starch was formedin the youngest leaves but there was a marked decline with leafage.By contrast, when replicate sets of discs were floated on waterand illuminated, photosynthetic starch formation was similarin the differently aged leaves. Uptake of sugar, measured bydry weight increases and incorporation of [¹⁴C]sucrose, wasnot dependent on leaf age. The possibility that physiologicalchanges, relating to ageing and import/export status of theleaf, regulate the metabolism of sugar to starch was examined.Increasing retention of sugar in the minor veins is likely tobe a major factor. Invertase activities were measured and foundto be similar in the differently aged leaves. Respiration ratesdeclined with increasing leaf age. Speculations concerning changesin selective permeability of the chloroplast membrane are alsodiscussed.     

Nitrate Supply and the Biophysics of Leaf Growth in Salix viminalis

The influence of nitrogen on leaf area development and the biophysicsof leaf growth was studied using clonal plants of the shrubwillow, Salix viminalis grown with either optimal (High N) orsub-optimal (Low N) supplies of nitrate. Leaf growth rate andfinal leaf size were reduced in the sub-optimal treatment andthe data suggest that in young rapidly growing leaves, thiswas primarily due to changes in cell wall properties, sincecell wall extensibility (% plasticity) was reduced in the LowN plants. The biophysical regulation of leaf cell expansion also differedwith nitrogen treatment as leaves aged. In the High N leaves,leaf cell turgor pressure (P) increased with age whilst in theLow N leaves P declined with age, again suggesting that foryoung leaves, cell wall plasticity limited expansion in theLow N plants. Measurements of cell wall properties showed thatcell wall elasticity (%E) was not influenced by nitrogen treatmentand remained constant regardless of leaf age. Key words: Salix, cell wall extensibility, nitrogen nutrition, biophysics of leaf growth     

The Role of Roots in Control of Bean Shoot Growth

Restriction of root growth by growing bean plants (Phaseolusvulgaris L.) in very small pots led to the development of dwarfplants. The leaves of those plants were smaller and their internodesshorter than those of control plants which were grown in largerpots and had developed a more extensive root system. A largequantity of starch—much more than in control plants —accumulated in the leaves and shoots of the dwarf plants. Increasingthe amount of minerals which was supplied to the roots, enhancedleaf growth of the control plants but failed to affect the dwarfones, in spite of the fact that in both cases the treatmentincreased the content of N, P and K in all the plant organs.The leaf water content was similar in both treatments, but theleaf water potential was higher in the dwarf plants. Exogenousapplication of gibberellic acid (GA₃) to the dwarf plants overcamethe reduction of stem growth completely, and that of the leavespartially. Application of the cytokinin, benzyladenine (BA)did not affect stem growth, but increased that of the primaryleaves. A combined supply of GA₂ + BA restored completely thegrowth of the stem and the primary leaves, and partially thatof the trifoliate leaves. It is concluded that a limited rootsystem restricts shoot growth through an hormonal system inwhich at least gibberellins and cytokinins are involved, andthat the dwarfing is not a consequence of mineral or assimilatedeficiency, or due to water stress. Phaseolus vulgaris L., leaf growth, stem growth, root restriction, gibberellic acid, benzyladenine, cytokinin     

Effects of leaf age, nitrogen nutrition and photon flux density on the organization of the photosynthetic apparatus in leaves of a vine (Ipomoea tricolor Cav.) grown horizontally to avoid mutual shading of leaves

Effects of leaf age, nitrogen nutrition and photon flux density (PFD) on the organization of the photosynthetic apparatus in leaves were investigated in a vine, Ipomoea tricolor Cav., which was grown horizontally so as to avoid mutual shading of leaves. The plants were grown hydroponically at two nitrate levels under two growth light treatments. For one group of the plants, leaves were exposed to full sunlight. For another group, respective leaves were artificially shaded in a manner that simulated changes in the light gradient with the development of an erect herbaceous canopy: old leaves were placed under progressively shadier conditions with growth of the plants (canopy-type shading). In all the treatments, chlorophyll (Chl) content gradually decreased with leaf age. Photosystem I (PSI) per Chl was constant, independent of leaf age, nitrogen nutrition and/or PFD. Photosystem II (PSII) and cytochrome / per Chl, and Chl a/b ratio were independent of leaf age and/or nitrogen nutrition but decreased with the decrease in growth PFD. Ribulose-1,5-bisphosphate carboxylase (EC 4.1.1.39, RuBPCase) per Chl steeply decreased with decrease in PFD. When leaves grown at the same PFD were compared, RuBPCase/Chl was lower in the plants grown under lower nitrogen availability and also decreased with leaf age in the plants grown without shading. These decreases were attributed to the curvilinear relationship between RuBPCase and Chl in leaves grown at full sunlight, that was independent of nitrogen availability and leaf age. From these results, it is concluded that the composition of the photosynthetic apparatus is independent of leaf age but changes depending on the light environment and total amount of photosynthetic components of the leaf.Abbreviations Chl chlorophyll - cyt f cytochrome f - PFD photon flux density - RuBPCase ribulose-1,5-bisphosphate carboxylase The author thanks Drs. K. Sonoike, Y. Kashino, K. Okada, H. Hatanaka, Y. Suzuki and A. Aoyama for technical advise. The author also thanks Drs. I. Terashima, A. Makino (Tohoku University, Sendai, Japan), Dr. J.R. Evans (Research School of Biological Sciences, Australian National University, Canberra) and Prof. A. Watanabe for valuable suggestions.     

The Effect of Temperature during Growth on the Subsequent Rate of Photosynthesis in Leaves of Tall Fescue (Festuca arundinacea Schreb)

Two experiments are reported in which young plants of tall fescuewere grown in temperature regimes of 20 °C day/15 °Cnight or 10 °C day/5 °C night until the fourth leafon the main stem was fully expanded. These temperature regimeswere then either changed over for individual plants or continuedunchanged up to the seven-leaf stage. Photosynthesis and respirationrates were determined in the fourth and subsequent leaves andalso in ageing leaves, using an infra-red gas analyser in anopen system and at temperatures of 10 and 20 °C in one and10, 15, 20, and 25 °C in the other experiment. Rates of apparent photosynthesis per unit leaf area in fullyexpanded leaves differed little as a result of previous treatmentand were not greatly affected by temperature during measurement.However, the specific leaf area and the rate of apparent photosynthesisper unit dry weight were higher in plants grown at the hightemperature. Leaves from the high-temperature regime had a higheroptimum temperature for apparent photosynthesis, a shorter life,and a lower respiration rate at any one temperature of measurementthan did leaves from the low-temperature regime. After transfer from one temperature regime to the other, therate of apparent photosynthesis of the next leaf to become fullyexpanded was higher in plants transferred from low to high temperatureand lower in plants transferred from high to low than in plantsremaining in either temperature regime; the leaves which subsequentlyexpanded had rates similar to those of unchanged plants. Inleaves which were fully expanded at the time of transfer, therate of apparent photosynthesis rose after transfer to the high-temperatureregime and fell after transfer to the low-temperature regime. These results are discussed in relation to growth-analysis datafrom other plants grown in the same conditions.     

Studies on the Expansion of the Leaf Surface: IV. THE CARBON AND PHOSPHORUS ECONOMY OF A LEAF

The fixation, utilization, and translocation of carbon and thenet import and export of phosphorus by three leaves of Cucumissativus over the course of their lives were measured in a controlledenvironment. The rate of photosynthesis of a leaf followed a regular dailypattern, rising to a maximum during the first 2 hrs. of thelight period and subsequently falling. Dark respiration wasusually highest at the beginning of the dark period and fellthroughout it. The daily rate of photosynthesis per unit areaof a leaf fell during its later life partly as a result of shadingby upper leaves and also because of an independent age factor.The rate of dark respiration per unit area was high in veryyoung leaves but fell rapidly with age. The amount of phosphorus in each leaf reached a maximum beforethe leaf had reached its maximum dry weight. There was thensubstantial net loss of phosphorus from the leaf. The changing function of each leaf as a sink or source of carbohydrateand mineral nutrients was determined. Four stages were recognized:(1) early development from inception until some time after unfolding,when the leaf was dependent upon imported carbohydate; (2) aperiod of rapid expansion, associated with a high rate of uptakeor mineral nutrients, during which translocation of assimilatedcarbon from the leaf was most rapid; (3) a time of decliningrates of growth, photosynthesis and export of carbon, associatedwith substantial loss of phosphorus; (4) finally, a short sensescentphase with net loss of CO₂, terminating in the death of theleaf.     

The Dynamics of Carbon Supply from Leaves of Barley Plants Grown in Long or Short Days

The role of the mature leaf in supplying carbon for growth inother parts of the plant was examined using a steady-rate ¹⁴CO₂labelling technique. The pattern of events occurring in theleaf during one complete 24 h cycle was compared in plants grownin, and adapted to long and short photoperiods. The rates ofleaf photosynthesis, night respiration and daytime loss of carbonfrom the growing regions of the plant Were similar in long orshort photoperiods. As a percentage of the total carbon fixedduring the photoperiod, total respiration was c. 50% for shortday plants but only 25% for long day plants. Thirty to forty per cent of the carbon fixed during the photoperiodwas retained in the leaf for export during darkness—therest was exported immediately. In leaves of short day plantssucrose and starch were the main form of the stored carbon.By the end of the dark period these compounds had been almostcompletely depleted. In leaves of long day plants there weremuch larger basal levels of sucrose and starch, upon which thediurnal variations were superimposed. These leaves also accumulatedfructosans. The delay in starch remobilization previously foundin leaves of short day plants was also evident in leaves oflong day plants even though large concentrations of sucroseand fructosans were present This suggests the presence of distinctpools of sucrose in the leaf.     

Ineffectiveness of cytokinin-induced chlorophyll retention in hypostomatous leaf discs

Kinetin retarded the decrease in chlorophyll content in leafdiscs from 5 species of plants with amphistomatous leaves, wherethe upper surface was exposed to air, but not in Rumex acetosera.When leaf discs were floated so that the lower surface was exposed,the effect of kinetin was less evident. Kinetin also stimulatedtranspiration in leaf discs from Nicotiana tabacum (amphistomatous),but not in leaf discs from Paederia chinensis (hypostomatous).Nor kinetin did retard chlorophyll breakdown in this specieswhen leaf discs were floated so that the stomatal surface wasin contact with the solution. The ineffectiveness of cytokininsin chlorophyll retention in leaf discs from hypostomatous leaveswas not due to reduced uptake of benzylaminopurine-¹⁴C. Chlorophyll retention was severely inhibited by coating theleaf surface with vaseline either in the presence or absenceof kinetin. Leaf discs floated on a solution exposed to CO₂-lessair retained more chlorophyll than those in normal air. Thereis thus a close relationship between stomatal opening (as measuredby stimulation of transpiration) and chlorophyll retention,as influenced by cytokinins. It is suggested that cytokinin-induced chlorophyll retentionand odier effects on leaf tissues could be mediated throughits effects on stomatal opening. (Received January 22, 1976; )     

Content and Distribution of Erythroxylum coca Leaf Alkaloids

Fully expanded leaves of Erythroxylum coca var. coca Lam. (Erythroxylaceae)35-70-d-old were harvested from 21-month-old plants grown undergreenhouse conditions. Each harvested leaf was placed on a plexiglasssilhouette according to its dimensions and divided into fourprimary sections (petiole, base, mid and anterior) and threesubsections (lamina periphery, false mid-rib, and true mid-rib)to determine the distribution and content of hygrine, cuscohygrine,trans-cinnamoylcocaine, cis -cinnamoylcocaine, tropacocaine,tropinone, methyl ecgonine and cocaine. The leaf sub-sectionsand petiole were extracted and analysed for alkaloid content(%) by HPLC (cocaine alkaloid only) gas chromatography and GC/MS.Cocaine, methyl ecgonine and hygrine were highest in the laminaperiphery with a content of 0·48, 0·46 and 0·32%,respectively. Trans-cinnamoylcocaine was pre-eminent of thecinnamoylcocaines and was most abundant in the petiole at acontent of 0·24%. Cis-cinnamoylcocaine, tropinone, cuscohygrineand tropacocaine were ubiquitously distributed throughout theleaf where the average contents were 0·14, 0·004,0·16 and 0·04%, respectively.Copyright 1995, 1999Academic Press Alkaloids, Erythroxylum coca var. coca, E. coca, leaf alkaloids, petiole alkaloids, cocaine, cuscohygrine, methyl ecgonine, hygrine, tropinone, tropacocaine, trans-cinnamoylcocaine, cis-cinnamoylcocaine     

Use of the Pressure Chamber in Membrane Damage Studies

The ability to detect membrane damage in plant leaves by thepressure chamber technique was evaluated. Membrane damage wasinduced by freezing and thawing, absorption of the host-specifictoxin victorin, ozonation, inoculation with Helminthosporiummaydis, and spraying with the phytotoxin, fusicoccin. The pressure-volumerelationships, i.e. the volume of sap expressed from a leafwith incremental increases in pressure, were compared in leaveswith intact or damaged membranes. Where membrane damage waswidespread throughout the leaf, sap was expressed at pressuresas low as one tenth of those needed in leaves with intact membranesand at low pressures the amount of water expressed from leaveswith damaged membranes was up to 10 times that from leaves withintact membranes. Further, the pressure-volume curves of healthyleaves became linear when the leaf turgor potential was reducedto zero, but were non-linear in leaves with damaged membranes.Ozone treatment, inoculation with H. maydis, or spraying withfusicoccin damaged only a proportion of the leaf cells, andpressure-volume relationships more nearly resembled those obtainedwith healthy leaves rather than those obtained with freezingand thawing and absorption of victorin. It is concluded that the pressure chamber can be used to observemembrane integrity, but its ability to detect damage may belimited to conditions in which the majority of cells in theleaf are damaged.     

The Assessment of Speckled Leaf Blotch in Winter Wheat in New Zealand

Disease symptoms of speckled leaf blotch (Mycosphaerella graminicola)were first observed in early August, 86 d after sowing a winterwheat field trial in New Zealand. Disease hastened senescenceof all leaves, delayed the expansion of new leaves during springand reduced the maximum size of some later-formed leaves. Theabsolute green leaf area was reduced by disease through to fullsenescence of leaves, which occurred one week earlier than inhealthy plants. Disease indices calculated from the data onpercentage diseased area alone indicated a decline in diseaseduring rapid growth of the plants in spring as the temperatureincreased. However, this apparent decline was generated by themethod of calculation and was not evident when effects on leafarea were also considered. Senescence induced by the pathogenwas an important aspect of the disease syndrome. Mycosphaerella graminicola, Septoria tritici, speckled leaf blotch, winter wheat, yield loss     

Effects of High Air and Soil Temperature Stress on Growth and Tuberization in Solanum tuberosum

Potato plants (Solanum tuberosum L.) were grown at differentair and soil temperatures to determine the effects of high-temperaturestress on root, tuber, and shoot growth. Cooling the soil (17–27C) at high air temperatures (30–40 C) relieved noneof the visible symptoms of heat stress on shoot growth; norwas the degree of induction to tuberize in leaves increased,as reflected in tuberization of leaf-bud cuttings. Heating thesoil (27–35 C) at cool (17–27 C) air temperatureshad no apparent detrimental effect on shoot growth or inductionof leaves to tuberize. However, in each case hot soil largelyeliminated tuber development. In one experiment stolons grewup out of the hot soil and formed aerial tubers upon reachingthe cool air. When leaf-bud cuttings from induced plants wereused as a model system, high soil temperatures inhibited tuberdevelopment from the buried leaf buds, in the absence of anyroot growth. Apparently the induction of leaves to tuberizeis affected principally by air rather than soil temperature,but expression of the signal to tuberize can be blocked by highsoil temperature. Solanum tuberosum L., potato, temperature stress, soil temperature, tuberization     

Relative Abundance of Nickel in the Leaf Epidermis of Eight Hyperaccumulators: Evidence that the Metal is Excluded from both Guard Cells and Trichomes

Scanning electron microscopy combined with X-ray microanalysiswas used to localize the sites of nickel accumulation on theleaf epidermis of eight nickel accumulators grown on ultramaficsoils in Greece. In all species, nickel was excluded from guardcells, and in species possessing hairy leaves (seven out ofeight) nickel was excluded from the hairs. In some species themetal was present in subsidiary cells, yet at low levels, whilethe sites of higher accumulation were epidermal cells away fromstomata. Results indicate that nickel is not compatible withthe functions and development of certain epidermal cell types.Copyright 2000 Annals of Botany Company Brassicaceae, epidermis, metal hyperaccumulators, microanalysis, serpentine plants     

A Positive Root-sourced Signal as an Indicator of Soil Drying in Apple, Malus x domestica Borkh.

Plants raised from shoot-tip cultures of apple (Malus x domesticaBorkh ) were grown in divided pots so that approximately halfthe root system could be exposed to soil drying whilst the remainderwas well watered. Water was withheld from half the root systemin this way for 24 d. During this time, the daily incrementin leaf area declined to 65% of that of control plants, whichhad all their roots in well watered soil Both leaf expansionand, more particularly, leaf initiation were inhibited. Waterloss per plant also fell to 70%% of the rate exhibited by controlplants. No significant differences were detected between theleaf water status of the partially dried plants and that ofthe well watered ones After this drying period, the half-dried plants were treatedduring the night of the 24th day as follows; one group had itsroots in dry soil rewatered, another had them excised, and athird continued half dry Both the rewatered group and the groupwith roots excised showed significant recoveries in leaf growthrates compared with the group kept half dry. The inhibition of leaf growth by drying soil is discussed withregard to root-to-shoot signalling The alleviation of this inhibitionby the excision of roots in dry soil, is taken as evidence ofa positive inhibitor, produced by drying roots, influencingshoot growth. Key words: Leaf growth, soil drying, root signals, Malus x domestica     

Effects of leaf age,nitrogen nutrition and photon flux density on the distribution of nitrogen among leaves of a vine (Ipomoea tricolor Cav.) grown horizontally to avoid mutual shading of leaves

Effects of leaf age, nitrogen nutrition and photon flux density (PFD) on the distribution of nitrogen among leaves were investigated in a vine, Ipomoea tricolor Cav., which had been grown horizontally so as to avoid mutual shading of leaves. The nitrogen content was highest in newly developed young leaves and decreased with age of leaves in plants grown at low nitrate concentrations and with all leaves exposed to full sunlight. Thus, a distinct gradient of leaf nitrogen content was formed along the gradient of leaf age. However, no gradient of leaf nitrogen content was formed in plants grown at a high nitrate concentration. Effects of PFD on the distribution of nitrogen were examined by shading leaves in a manner that simulated changes in the light gradient of an erect herbaceous canopy (i.e., where old leaves were placed under increasingly darker conditions with growth of the canopy). This canopy-type shading steepened the gradient of leaf nitrogen content in plants grown at a low nitrogen supply, and created a gradient in plants grown at high concentrations of nitrate. The steeper the gradient of PFD, the larger the gradient of leaf nitrogen that was formed. When the gradient of shading was inverted, that is, younger leaves were subjected to increasingly heavier shade, while keeping the oldest leaves exposed to full sunlight, an inverted gradient of leaf nitrogen content was formed at high nitrate concentrations. The gradient of leaf nitrogen content generated either by advance of leaf age at low nitrogen availability, or by canopy-type shading, was comparable to those reported for the canopies of erect herbaceous plants. It is concluded that both leaf age and PFD have potential to cause the non-uniform distribution of leaf nitrogen. It is also shown that the contribution of leaf age increases with the decrease in nitrogen nutrition level.