Influence of Genome on Leaf Anatomy of Triticum and Aegilops

The leaf mesophyll of Triticum and Aegilops is constructed fromcells with one to ten arms. Volume of mesophyll cells per unitleaf area was larger in some monogenomic (A and B genome) plantsthan in polyploids, while leaf volume per unit leaf area wassmaller in the former than in the latter. Consequently, thecompactness of leaf blade is higher in these monogenomic plantsthan in the polyploids. D genome plants showed a much lowervolume of both mesophyll cells and leaf blade per unit leafarea, but the compactness of the leaf blade was generally higherthan in the polyploids. Mesophyll surface area per unit leaf area tended to be largerin the A and B genome than in the D genome and polyploid plants.Out of the polyploids, AB genome plants showed a larger mesophyllsurface area per unit leaf area as compared with AG and ABDgenome plants. Therefore, either the D or the G genome seemsto have the effect of decreasing the mesophyll surface areaper unit leaf area. A decrease of the compactness of leaf bladeand the mesophyll surface area per unit leaf area in the polyploidswas considered to be associated with the reduction of theirphysiological activities on the unit leaf area basis. Triticum, Aegilops, wheat, mesophyll surface area, leaf anatomy, genome, photosynthesis     

Variation in Mesophyll Cell Number and Size in Wheat Leaves

The numbers of mesophyll cells in wheat leaves were determinedin a variety of wheat species differing in ploidy level andin leaves from different positions on the wheat plant. Leafsize and mesophyll cell number are linearly related in bothcases but differences were observed in mesophyll cell numberper unit leaf area with changing leaf size. Where changes incell size are caused either by nuclear ploidy or leaf position,differences in mesophyll cell number per unit leaf are negativelycorrelated with mesophyll cell plan area. The decrease in cellsize with increasing leaf position also results in a greaternumber of chloroplasts per unit leaf area. These results arediscussed in relation to anatomical variation of the wheat leaf. Mesophyll cell, cell numbers, leaf size, Triticum     

Relationship between Leaf Structure and Gas Exchange in Wheat Leaves at Different Insertion Levels

Net photosynthesis rate (P_n), stomatal conductance to CO₂ andresidual conductance to CO₂ were measured in the last six leaves(the sixth or flag leaf and the preceding five leaves) of Triticumaestivum L. cv. Kolibri plants grown in Mediterranean conditions.Recently fully expanded leaves of well-watered plants were alwaysused. Measurements were made at saturating photosynthetic photonflux density, and at ambient CO₂ and O₂ levels. The specificleaf area, total organic nitrogen content, some anatomical characteristics,and other parameters, were measured on the same leaves usedfor gas exchange experiments. A progressive xeromorphic adaptation in the leaf structure wasobserved with increasing leaf insertion levels. Furthermore,mesophyll cell volume per unit leaf area (V_mes/A) decreasedby 52·6% from the first leaf to the flag leaf. Mesophyllcell area per unit leaf area also decreased, but only by 24·5%.However, nitrogen content per unit mesophyll cell volume increasedby 50·6% from the first leaf to the flag leaf. This increasecould be associated to an observed higher number of chloroplastcross-sections per mm² of mesophyll cell cross-sectional areain the flag leaf: values of 23000 in the first leaf and 48000in the flag leaf were obtained. P_n per unit leaf area remainedfairly constant at the different insertion levels: values of33·83±0·93 mg dm^–2 h^–1 and32·32±1·61 mg dm^–2 h^–1 wereobtained for the first leaf and the flag leaf, respectively.Residual conductance, however, decreased by 18·2% fromthe first leaf to the flag leaf. Stomatal conductance increasedby 41·7%. The steadiness in P_n per unit leaf area across the leaf insertionlevels could be mainly accounted for by an opposing effect betweena decrease in V_mes/A and a more closely packed arrangement ofphotosynthetic apparatus. Adaptative significance of structuralchanges with increasing leaf insertion levels and the steadinessin P_n per unit leaf area was studied. Key words: Photosynthesis, structure, wheat     

Leaf Characteristics and Net Gas Exchange of Diploid and Autotetraploid Citrus

The effect of tetraploidy on leaf characteristics and net gasexchange was studied in diploid (2x ) and autotetraploid (4x) ‘Valencia’ sweet orange (Citrus sinensis (L.)Osb.) and ‘Femminello’ lemon (Citrus limon (L.)Burm. f.) leaves. Comparisons between ploidy levels were madeunder high irradiance (I) in a growth chamber or low total Iin a glasshouse. Tetraploids of both species had thicker leaves,larger mesophyll cell volume and lower light transmittance thandiploids regardless of growth I. Mesophyll surface area perunit leaf area of 2x leaves was 5–15% greater than on4x leaves. Leaf thickness and mesophyll cell volume were greaterin high I leaves than low I leaves. In high I, average leafarea was similar for 2x and 4x leaves, whereas in low I it was30% greater in 4x than in 2x leaves. Nitrogen and chlorophyllconcentration per cell increased with ploidy level in both growthconditions. The ratio of chlorophyll a:b was 25% greater in2x than in 4x leaves. When net CO₂assimilation rate (A_CO2) wasbased on leaf area, 4x orange leaves had 24–35% lowerA_CO2than their diploids. There were no significant differencesin A_CO2between 2x and 4x orange or lemon leaves when expressedon a per cell basis. Overall, lower A_CO2per unit leaf area oftetraploids was related to increase in leaf thickness, largermesophyll cell volume, the decrease in mesophyll area exposedto internal air spaces, and the lower ratio between cell surfaceto cell volume. Such changes probably increased the resistanceto CO₂diffusion to the site of carboyxlation in the chloroplasts. Cell volume; chlorophyll; irradiance; leaf thickness; nitrogen; photosynthesis; ploidy; Citrus limon ; C. sinensis ; ‘Valencia’ sweet orange; ‘Femminello’ lemon     

Flag Leaf Photosynthesis of Triticum aestivum and Related Diploid and Tetraploid Species

Rates of net photosynthesis of the flag leaves of 15 genotypesof wheat and related species were measured throughout theirlife, using intact leaves on plants grown in the field. At thestage when rates were maximal, they were in general highestfor the diploid species, intermediate for the tetraploidspeciesand lowest for Triticum aestivum (means of 38, 32 and 28 mgCO₂ dm^–2 h^–1 respectively). Rates were stronglynegatively correlated with leaf area, leaf width and the meanplan area per mesophyll cell and positvely correlated with stomatalfrequency and number of veins per mm of leaf width. The differencesamong species in these attributes were mainly related to ploidylevel. It was not possible to determine the relative importanceof each anatomical feature, though the changes in stomatal frequencyhad only slight effects on stomatal conductance and the observeddifferences in rates of photosynthesis were much greater thanwould be expected from those in stomatal conductance alone. There was genetic variation in rates of light dependent oxygenevolution of isolated protoplasts and intact chloroplasts butno difference attributable to ploidy. The mean rate, 91 µmolO₂ mg^–1 chlorophyll h^–1, equivalent to 3.9 mg CO₂mg^-1chlorophyll h^-1 was considerably less than the rate of photosynthesisin comparable intact leaves, which was 7.2 mg CO₂ mg^–1chlorophyll h^–1. The total above-ground dry matter yields were least for thewild diploids T. urartu and T. thauodar and the wild tetraploidT. dicoccoides, but the other wild diploids produced as muchdry matter as the hexaploids. The prospects of exploiting differences in photosynthetic ratein the breeding of higher yielding varieties are discussed. Triticum aestivum L., wheat, Aegilops spp, photosynthesis, stomatal conductance, stomatal frequency, polyploidy     

Functional consequences of stenophylly for leaf productivity: comparison of the anatomy and physiology of a rheophyte, Farfugium japonicum var. luchuence, and a related non-rheophyte, F. japonicum (Asteraceae)

We investigated the anatomical and physiological characteristics of stenophyllous leaves of a rheophyte, Farfugium japonicum var. luchuence, and sun and shade leaves of a non-rheophyte, F. japonicum, comparing three different populations from coastal, forest floor, and riparian habitats. Light adaptation resulted in smaller leaves, and riparian adaptation resulted in narrower leaves (stenophylly). The light-saturated rate of photosynthesis (P _max) per unit leaf area corresponded to the light availability of the habitat. Irrespective of leaf size, the P _max per unit leaf mass was similar for sun and shade leaves. However, the P _max per mass of stenophyllous leaves was significantly lower than that of sun and shade leaves. This was because the number and size of mesophyll cells were greater than that required for intercellular CO₂ diffusion, which resulted in a larger leaf mass per unit leaf area. Higher cell density increases contact between mesophyll cells and enhances leaf toughness. Stenophyllous leaves of the rheophyte are frequently exposed to a strong water flow when the water level rises, suggesting a mechanical constraint caused by physical stress.     

Effects of Constant and Variable Nitrogen Supply on Sunflower (Helianthus annuus L.) Leaf Cell Number and Size

The effects of nitrogen (N) availability on cell number andcell size, and the contribution of these determinants to thefinal area of fully expanded leaves of sunflower (Helianthusannuus L.) were investigated in glasshouse experiments. Plantswere given a high (N =315 ppm) or low (N=21 ppm) N supply andwere transferred between N levels at different developmentalstages (5 to 60% of final size) of target leaves. The dynamicsof cell number in unemerged (< 0.01 m in length) leaves ofplants growing at high and low levels of N supply were alsofollowed. Maximum leaf area (LA_max) was strongly (up to two-fold)and significantly modified by N availability and the timingof transfer between N supplies, through effects on leaf expansionrate. Rate of cell production was significantly (P<0.05)reduced in unemerged target leaves under N stress, but therewas no evidence of a change in primordium size or in the durationof the leaf differentiation–emergence phase. In fullyexpanded leaves, number of cells per leaf (N_cell), leaf areaper cell (LA_cell) and cell area (A_cell) were significantly reducedby N stress. WhileLA_cell and A_cellresponded to changeover treatmentsirrespective of leaf size, significant (P<0.05) changes inN_cellonly occurred when the changeover occurred before the leafreached approx. 10% of LA_max. There were no differential effectsof N on numbers of epidermal vs. mesophyll cells. The resultsshow that the effects of N on leaf size are largely due to effectson cell production in the unemerged leaf and on both cell productionand expansion during the first phase of expansion of the emergedleaf. During the rest of the expansion period N mainly affectsthe expansion of existing cells. Cell area plasticity permitteda response to changes in N supply even at advanced stages ofleaf expansion. Increased cell expansion can compensate forlow N_cellif N stress is relieved early in the expansion of emergedleaves, but in later phases N_cellsets a limit to this response.Copyright 1999 Annals of Botany Company Helianthus annuus, leaf expansion, leaf cell number, leaf cell size, nitrogen, leaf growth, sunflower.     

Dry Matter Yields and Photosynthetic Rates of Diploid and Hexaploid Triticum Species

The above ground dry matter yields of two wild diploid Triticumspecies averaged 76 per cent of that of two hexaploid breadwheat varieties in field trials carried out over five years.A cultivated diploid species, T. monococcum, gave similar drymatter yields to the bread wheat varieties but had a longergrowth cycle. The flag leaves of wild diploid species had higher rates ofphotosynthesis than those of the bread wheat varieties bothwhen expressed per unit area of leaf or per unit weight of chlorophyll.Photosynthetic rates of other organs, expressed per unit weightof chlorophyll were also greater for the wild diploids thanfor hexaploids. For snoots at the stage when their flag leaveswere fully expanded, the investment in photosynthetic machinery,as measured by chlorophyll concentration, was less in the twowild diploids than in the hexaploids. This compensated for thehigh photosynthetic rate of the former, such that the specificgrowth rates, assessed by carbon-14 fixation per unit shootdry matter, were similar. Triticum spp., wheat, dry matter yield, photosynthesis, carbon-14, ploidy     

Changes in Size and Number of Mesophyll Cells, Nitrogen Content and Photosynthesis with Leaf Order in Brassica spp.

Numbers of mesophyll cells per unit leaf area decreased progressivelyfrom an upper leaf with a width of 3 cm towards the lower leaves.Enlargement of mesophyll cell size with leaf order accountedfor an increase or maintenance of mesophyll cell surface areaper unit leaf area. Increase of photosynthetic rates was correlatedwith increases of mesophyll cell surface area and nitrogen contentper unit leaf area. Therefore, in spite of an increase in cellsurface area to volume ratio with increase of mesophyll cellsize, it appears that increase of mesophyll cell surface areaand nitrogen content per unit leaf area enables a high rateof photosynthesis to be maintained. Brassica, photosynthesis, mesophyll surface area, nitrogen content, cell size, mesophyll resistance, leaf age     

Leaf expansion and carbon assimilation in cotton leaves grown at two photosynthetic photon flux densities

Increasing photosynthetic photon flux density (PPFD) received during development from 5.5 to 31.2 mol m^-2 d^-1 resulted in greater leaf and mesophyll cell surface areas in cotton (Gossypium hirsutum L.). The relationships between the amounts of these surface areas and potential CO₂ assimilation by these leaves were evaluated. Leaf area (epidermal surface area of one side of a leaf), mesophyll cell surface area, and net rate of CO₂ uptake (P_n) were measured from the time leaves first unfolded until P., was substantially reduced. At the higher PPFD, leaf and mesophyll surface areas increased more rapidly during expansion, and P_n per unit leaf area was greater than at the lower PPFD. Although leaves at the higher PPFD reached the maximum P., per unit mesophyll cell surface area 4 to 5 days earlier than leaves at the lower PPFD, the maxima for these P., were similar. Leaves grown at the higher PPFD had the potential to assimilate 2.2, 3.5, or 5.8 times the amount of CO₂ as leaves from the lower PPFD when P., was expressed per unit mesophyll surface, per unit leaf surface, or per whole leaf, respectively. Greater and earlier development of both P., and mesophyll cell surface area at higher PPFD apparently had a compounding effect on the potential for carbon assimilation by a leaf.     

Genome-Dependent Factors in the Development of Leaf Phototrophic Tissues in Diploid and Alloploid Wheat Species

Growth and mesostructure of the photosynthetic apparatus were studied in leaves of ten Triticum L. species. Plants with the A^u genome were shown to develop larger leaf assimilation areas due to expanding areas of individual leaves and an increase in the absolute growth rate. Leaf and mesophyll thickness and mesophyll cell size decreased in the G-genome species. Leaf compactness, which depended on cell size and number per unit leaf area and leaf folding, determined the specific patterns of internal leaf organization in wheat species with diverse genotypes. These patterns did not affect cell plastid-to-cytoplasm ratio as shown by the stable indices of cell surface area/cell volume, cell surface area per chloroplast, and cell volume per chloroplast. The structural indices of leaf phototrophic tissues, mesophyll density, and mesophyll CO₂ conductance in alloploids, as compared to diploid species, depended on both ploidy and genome constitution.     

The Dynamics of Leaf Growth and Photosynthetic Capacity in Capsicum frutescens L.

In Capsicum frutescens L. cv. California Wonder the specificleaf weight (dry weight per unit laminar area) at leaf unfoldingis three times higher in the eighth leaf than in the first leafproduced. Intermediate leaves exhibit a trend between the twoThe change in specific leaf weight during laminar expansionis greatest in leaf 1 and least (sometimes zero) in leaf 8.Large changes in specific leaf weight during laminar expansionare associated with a large degree of palisade cell expansion,while leaves showing smaller rates of change have less palisadecell expansion but cell division is more evident. At leaf unfoldingthe fraction I protein content per unit laminar area is higherin upper than in lower leaves. Ribulose diphosphate carboxylaseactivity per unit laminar area and ¹⁴CO₂ fixation per unit laminararea have a similar pattern of development in all leaves andshow no correlation with the changes in specific leaf weight.The peak of activity in all leaves occurs when the laminar areais 10 cm². These results are compared with previous data onlaminar expansion and are seen as in accord with current ideason leaf growth.     

Changes of Anatomical Features, Photosynthesis and Ribulose Bisphosphate Carboxylase-Oxygenase Content of Mango Leaves

Changes in anatomical and physiological features, includingchanges in amount per unit area of anthocyanin and chlorophyll,in leaves of seedling mango (Mangifera indica L. cv. Irwin)trees were determined to understand what controls the rate ofphotosynthesis (Pn) at various stages of development. The youngleaves of seedling trees contained high concentrations of anthocyanin.During enlargement of leaves, the disappearance of anthocyaninand the accumulation of chlorophyll occurred concomitantly;the anthocyanin content began to decrease markedly once theleaf area had reached a maximum. During the early period ofleaf development, the thickness of mesophyll tissue decreasedtemporarily, but when the length of the leaf reached half thatof a mature leaf, the mesophyll began to thicken again. Smallstarch grains appeared in the chloroplasts of the young leavesand chloroplast nucleoids (ct-nuclei) were distributed throughoutthe chloroplasts. When leaves matured, ct-nuclei were displacedto the periphery of chloroplasts because of the accumulationof large starch grains. Compared with young leaves, green andmature leaves contained greater concentrations of ribulose bisphosphatecarboxylase-oxygenase (RuBisCO) protein. The results of immunocytochemicalexamination of RuBisCO under the light microscope reflectedthe results of electrophoresis measurements of RuBisCO. Pn waslow during the chocolate-coloured stage of early leaf development.In green and mature leaves Pn was higher; the average Pn was7·6 mg CO₂ dm^-2 h^-1 under light at intensities above500 µmol m^-2 s^-1.Copyright 1995, 1999 Academic Press Mangifera indica L., mango leaf, chloroplast nucleoids, chloroplast ultrastructure, starch accumulation, anthocyanin, chlorophyll, DAPI staining, SDS-PAGE, immunocytochemical technique     

Leaf anatomy and light acclimation in woody seedlings after gap formation in a cool-temperate deciduous forest

The photosynthetic light acclimation of fully expanded leaves of tree seedlings in response to gap formation was studied with respect to anatomical and photosynthetic characteristics in a natural cool-temperate deciduous forest. Eight woody species of different functional groups were used; two species each from mid-successional canopy species (Kalopanax pictus and Magnolia obovata), from late-successional canopy species (Quercus crispula and Acer mono), from sub-canopy species (Acer japonicum and Fraxinus lanuginosa) and from vine species (Schizophragma hydrangeoides and Hydrangea petiolaris). The light-saturated rate of photosynthesis (P _max) increased significantly after gap formation in six species other than vine species. Shade leaves of K. pictus, M. obovata and Q. crispula had vacant spaces along cell walls in mesophyll cells, where chloroplasts were absent. The vacant space was filled after the gap formation by increased chloroplast volume, which in turn increased P _max. In two Acer species, an increase in the area of mesophyll cells facing the intercellular space enabled the leaves to increase P _max after maturation. The two vine species did not significantly change their anatomical traits. Although the response and the mechanism of acclimation to light improvement varied from species to species, the increase in the area of chloroplast surface facing the intercellular space per unit leaf area accounted for most of the increase in P _max, demonstrating the importance of leaf anatomy in increasing P _max.     

The Structure of the Mesophyll of Flag Leaves in Three Triticum Species

Flag leaves of Triticum urartu, T. monococcum and T. aestivumcv. Professeur Marchal were examined by light and electron microscopyand by separating cells to determine whether differences inleaf anatomy could be related to known differences in theirlight-saturated rates of photosynthesis. Mesophyll cells fromthe three species were lobed and orientated with their longaxis parallel to the veins. The longest, most-lobed cells flankedthe sclerenchyma associated with the veins. Mean cell dimensionswere greatest in Professeur Marchal, but there was no significantdifference in the ratio of the mesophyll cell surface area tocell volume amongst the three species. Flag leaves of T. urartushowed the highest rates of photosynthesis and were also thethickest, with closely-spaced veins from which many of the mesophyllcells radiated. These flag leaves also had significantly more(21.9 per cent) air-filled space, and the highest ratio (15.2)of mesophyll cell surface exposed to this air-filled space perunit leaf area. Ways in which these anatomical characteristicsmay contribute to the higher rate of photosynthesis are discussed. Triticum urartu, Triticum monococcum, Triticum aestivum, flag leaves, morphology, mesophyll     

The Greenhouse Effect: Acclimation of Tomato Plants Growing in High CO2, Photosynthesis and Ribulose-1, 5-Bisphosphate Carboxylase Protein

Tomato plants were grown in solution culture in a controlledenvironment at 20 ?C with a 12 h photoperiod of 400 µmolquanta m^–2 s^–1 PAR with either normal ambient CO₂,approximately 340 vpm, or with 1000 vpm CO₂. The short- andlong-term effects of CO₂ enrichment on photosynthesis were determinedtogether with the levels of ribulose-1, 5-bisphosphate carboxylase(RuBPco) E.C. 4.1.1.39 [EC] protein and activity throughout leafdevelopment of the unshaded 5th leaf above the cotyledons. Thehigh CO₂ concentration during growth did not appreciably affectthe rate of leaf expansion or final leaf area but did increasethe fresh weight per unit area of leaf. With short-term CO₂enrichment, i.e. only during the photosynthesis measurements,the light-saturated photosynthetic rate (P_max) of young leavesdid not increase while those reaching full expansion more thandoubled their net rate of CO₂ fixation. However, with longerterm CO₂ enrichment, i.e. growing the crop in high CO₂, theplants did not maintain this photosynthetic gain. While theCO₂ concentration during growth did not affect the peak in P_maxmeasured in 300 vpm CO₂ or P_max in 1000 vpm CO₂, RuBPco proteinor its activity, the subsequent ontogenetic decline in theseparameters was greatly accelerated by the high CO₂ treatment.Compared with plants grown in normal ambient CO₂ the high CO₂grown leaves, when almost fully expanded, contained only approximatelyhalf as much RuBPco protein and P_max in 300 vpm CO₂ and P_maxin1000 vpm CO₂ were similarly reduced. The loss of RuBPco proteinmay be a major factor associated with the accelerated fall inP_max since it was close to that predicted from the amount andkinetics of RuBPco assuming RuBP saturation. In the oldest leavesexamined grown in high CO₂ additional factors may be limitingphotosynthesis since RuBPco kinetics marginally overestimatedP_max in 300 vpm CO₂ and the initial slope of photosynthesisin response to intercellular CO₂ was also less than expectedfrom the extractable RuBPco. Key words: Lycopersicon esculentum (Mill.) cv. Findon Cross, CO₂ enrichment, acclimation to high CO₂, photosynthesis, RuBPco protein and activity     

Does the photosynthetic light‐acclimation need change in leaf anatomy?

There is a strong correlation between leaf thickness and the light‐saturated rate of photosynthesis per unit leaf area (P_max). However, when leaves are exposed to higher light intensities after maturation, P_max often increases without increasing leaf thickness. To elucidate the mechanism with which mature leaves increase P_max, the change in anatomical and physiological characteristics of mature leaves of Chenopodium album, which was transferred from low to high light condition, were examined. When compared with leaves subjected to low light continuously (LL leaves), the leaves transferred from low to high light (LH leaves) significantly increased P_max. The transfer also increased the area of chloroplasts facing the intercellular space (S_c) and maintained a strong correlation between P_max and S_c. The mesophyll cells of LL leaves had open spaces along cell walls where chloroplasts were absent, which enabled the leaves to increase P_max when they were exposed to high light (LH). However, the LH leaves were not thick enough to allow further increase in P_max to the level in HH leaves. Thus leaf thickness determines an upper limit of P_max of leaves subjected to a change from low to high light conditions. Shade leaves would only increase P_max when they have open space to accommodate chloroplasts which elongate after light conditions improve.     

Photosynthesis by Flag Leaves of Wheat in Relation to Protein, Ribulose Bisphosphate Carboxylase Activity and Nitrogen Supply

The effects of nitrate supply on the composition (cell numbers,protein and chlorophyll contents) of flag leaves of winter wheatgrown with two amounts of N fertilizer and of spring wheat grownin the glasshouse under controlled nitrate supply are describedand related to photosynthesis. Nitrogen deficiency decreasedthe size of leaves, mainly by reducing cell number and, to asmaller extent, by decreasing cell volume. Protein content perunit leaf area, per cell and per unit cell volume was largerwith abundant N. Total soluble protein, ribulose bisphosphatecarboxylase-oxygenase (RuBPc-o) protein and chlorophyll changedin proportion irrespective of nitrogen supply and leaf age.Photosynthesis per unit area of flag leaf and carboxylationefficiency in both winter and spring wheat were proportionalto the amount of total soluble protein up to 7.0 g m^–2and to the amount of RuBPc-o protein up to 4.0 g m^–2.However, photosynthesis did not increase in proportion to theamount of total soluble or RuBPc-o protein above these amounts.In young leaves with a high protein content the measured ratesof photosynthesis were lower than expected from the amount andactivity of RuBPc-o. Carboxylation per unit of RuBPc-o protein,measured in vitro, was slightly greater in N-deficient leavesof winter wheat but not of spring wheat. RuBPc-o activity perunit of RuBPc-o protein was similar in winter and spring wheatleaves and remained approximately constant with age, but increasedin leaves showing advanced senescence. RuBPc-o protein fromN-deficient leaves migrated faster on polyacrylamide gels thanprotein from leaves with high N content. Regulation of the rateof photosynthesis in leaves and chloroplasts with a high proteincontent is discussed. The conductance of the cell to the fluxof CO₂ from intercellular spaces to RuBPc-o active sites iscalculated, from cell surface areas and CO₂ fluxes, to decreasethe CO₂ partial pressure at the active site by less than 0.8Pa at an internal CO₂ partial pressure of 34 Pa. Thus the decreasein partial pressure of CO₂ is insufficient to account for theinefficiency of RuBPc-o in vivo at high protein contents. Otherlimitations to the rate of photosynthesis are considered. Key words: Wheat, photosynthesis, nitrogen, ribulose, bisphosphate carboxylase     

Temporal and Spatial Development of the Cells of the Expanding First Leaf of Arabidopsis thaliana (L.) Heynh

The three-dimensional quantitative leaf anatomy in developingyoung (9–22 d) first leaves of wild type Arabidopsis thalianacv. Landsberg erecta from mitosis through cell and leaf expansionto the cessation of lamina growth has been studied. The domainsof cell division, the relative proportion of the cell typespresent during development and the production of intercellularspace in the developing leaf have been determined by image analysisof entire leaves sectioned in three planes. Mitotic activityoccurs throughout the youngest leaves prior to unfolding andcell expansion is initiated firstly at the leaf tip with a persistentzone of mitotic cells at the leaf base resulting in a gradientof development along the leaf axis, which persists in the olderleaves. Major anatomical changes which occur during the developmentare, a rapid increase in mesophyll volume, an increase in thevein network, and expansion of the intercellular spaces. Thepattern of cell expansion results in a 10-fold variation inmesophyll cell size in mature leaves. In the youngest leavesthe plan area of mesophyll cells varies between 100 µm²and 400 µm² whereas in mature leaves mesophyll cells rangein plan area from 800 µm² to 9500 µm². The volumesof mesophyll tissue and airspace under unit leaf area increase3-fold and 35-fold, respectively, during leaf expansion. Thevolume proportions of tissue types mesophyll:airspace:epiderrnal:vascularin the mature leaf are 61:26:12:1, respectively. This studyprovides comparative information for future identification andanalysis of leaf development mutants of Arabidopsis thaliana. Key words: Arabidopsis, quantitative leaf anatomy, leaf expansion, image analysis     

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

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