Development of the Fifth Leaf is Indicative for Whole Plant Performance at Low Temperature in Tomato

In the search for early-detectable selection criteria for growthat low temperature conditions in tomato, first the initiationand growth of individual leaves was analysed. Scanning electronmicroscopy revealed that the first four primordia had alreadydeveloped during the germination period at 25°C. The primordiumof the fifth leaf, however, was initiated after the transferof seedlings to the experimental conditions. The increase inlength of the first three leaves, and to a lesser extent ofthe fourth leaf, was considerably smaller in comparison withthat of later formed leaves. Moreover, the morphology of thefirst three to four leaves was deviant, whereas the others showedthe normal compound leaf architecture. All these results indicatedthat the fifth leaf was the earliest formed leaf with growthcharacteristics that might reflect the growth potential of thewhole plant. Development of the fifth leaf was tested as a marker for wholeplant growth. At three temperature, 18, 15 and 12°C, growthresponses of the fifth leaf were similar to that of whole plantsin four tomato genotypes: Line A, Line B, Premier and MXXIV-13.Significant differences in relative growth rate of dry weightof whole plants and fifth leaves (RGR_W)and of leaf area of thefifth leaves (RGR_LA between two fast growing and two slow growinggenotypes were found. No genotype by temperature interactionfor RGR_W and RGR_LA was found, indicating that the effect oftemperature decrease was similar for the four genotypes. The structure of the mature fifth leaf of one fast and one slowgrowing genotype, Line A and MXXIV-13, was analysed. For bothgenotypes, leaves were small and thick at low temperature, 12°C.The total number of epidermis and palisade parenchyma cellsper leaf was smaller but the size of the cells developed at12°C was larger than at 18°C. Consequently, the slowgrowth at 12°C was due to a low rate of cell division. Atboth temperatures, the fifth leaf to MXXIV-13 was smaller comparedto that of line A. Since the size of the cells were similar,the smaller leaf size was due to lower number of leaf cells. The results confirm the suitability of the growth, especiallyexpressed as RGR_LA , of the fifth leaf as a nondestructive marketfor vegetative development of tomato at low temperature. Growthdifferences between genotypes were mainly reflected by differencesin cell number of leaves, which might be correlated with geneticallydetermined differences in cell number of leaf primordia.Copyright1993, 1999 Academic Press Lycopersicon esculentum Mill. genotypes, plant growth, selection criteria, low temperature, leaf initiation, leaf development, RGR, leaf structure, cell expansion     

Some Effects of Temperature and Irradiance on Growth of the First Four Leaves of Wheat, Triticum aestivum

Plants of Heron wheat were grown at 20 and 15 °C and inquantum flux densities of 400 and 200 µmol m^–2 s^–1.At completion of expansion of the first or second leaf, plantswere transferred between temperatures and quantum flux densities.Final size and cell number were measured for each of the firstfour main-stem leaves. Leaf area was affected only slightlyby treatment and effects on leaf length and width were alsosmall. It was concluded that leaf extension rate, which waslower at the lower temperature and in the lower light regime,is inversely related to the duration of leaf expansion. Leafdry wt was higher for plants grown in high light and for plantsgrown at 15 °C; transfer treatments led to readjustmentswhereby dry wts of leaves expanded after transfer resembledthose of leaves on plants kept throughout in the post-transferconditions. Leaf cell number was not affected by treatment but mean drywt per cell was significantly greater in high light, and forthe first two leaves, at 15 °C. There was a major and highlysignificant effect of treatment on the ratio of dry: fresh wtper cell, this being larger for leaves in high light. Transfertreatments between light regimes led to rapid changes in expandingleaves as was found for leaf dry wt. It was concluded that theexpanding grass leaf is much less dependent on older leavesto provide the necessary materials for cell division and expansionthan is the dicotyledon leaf. It is suggested that the increasein cell dry wt in high light is associated with an increasein cell wall material which is under photomorphogenic control. Triticum aestivum, wheat, leaf growth, cell division, cell expansion, cell size     

The Effects of Position and Temperature on the Expansion of Leaves of Vicia faba L.

The growth in area of the first eight leaves of broad bean plantswas investigated in growth room experiments. Plants were grownat either 20 or 14 °C or transferred from 20 to 14 °C.Rates of leaf appearance and unfolding increased with temperature.The duration of growth of a leaf increased with leaf numberfor the first five leaves and then remained constant The meangrowth rate declined or remained constant with increasing leafnumber Durations of growth were shorter and growth rates largerat 20 °C than at 14 °C Plants responded immediatelyto the change in temperature Final areas of leaves which expandedafter transfer from 20 to 14 °C were larger than those grownat 20 °C Vicla faba L., broad bean, leaf expansion, temperature responses     

The Effect of Temperature on the Growth of Cocksfoot (Dactylis glomerata L.)

The growth of cocksfoot at 14°, 22°, and 26° C wasmeasured at weekly intervals over a period of six weeks. Initially,the relative growth rates increased with increase in temperature,but during the final three weeks they were little differentat all three temperatures. The reduction in relative growthrates with time at 22° and 26° were associated withincreases in size which were partly reflected by reductionsin the leaf-area ratios. It is also likely that at 26° changesin the photosynthetic capacity of leaves, perhaps associatedwith decreasing concentrations of mineral nutrients, contributedto the decreased relative growth rates. Leaf expansion and increase in cell numbers were estimated overtwo-day periods at temperatures ranging from 5° to 30°C. Leaf expansion increased with increase in temperature throughoutthis range; extrapolation suggested that it would cease at temperaturesbelow 3° C. The optimum temperature for cell division appearedto be between 20° and 25°C. Different physiological processes appeared to be involved inthe temperature responses of plants of different sizes and histories.With young plants these responses resulted in a large overalleffect of temperature on the growth rate; with older plantsof the same size there appeared to be several compensatory responsesso that variation in temperature over an apprecaible range hadlittle overall effect.     

The Effects of Temperature on Leaf Growth in Cyperus longus, a Temperate C4 Species

Plants of the C₄ sedge Cyperus longus L. were grown at 10, 20and 30 °C. An asymptotic growth curve, the Richards function,was fitted to growth data for successive leaves. The mean rateof leaf appearance was a linear function of temperature with0.014 leaves appearing per day for every 1 °C increase intemperature. The instantaneous relative rate of leaf extensionshowed a marked ontogenetic drift which was most rapid at 30°C and slowest at 10 °C. The mean absolute extensionrate for foliage had a temperature coefficient of 0.16 cm d^–1° C^–1 in the range from 10 to 30 °C. The durationof leaf growth was independent of leaf number at 10 and 20 °Cbut increased linearly with leaf number at 30 °C. The smalldifferences in relative growth rate at the three temperaturesresulted in large differences in foliage area produced at theend of a 30 d growth period. The final foliage areas at 20 and10 °C were 51 and 9% respectively of that at 30 °C. Cyperus longus, temperature, leaf growth, Richards function, growth analysis     

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.     

The Growth and Development of Maize (Zea mays L.) at Five Temperatures

The objectives of this work were to measure growth and developmentrates over a range of temperatures and to identify processeswhich may limit vegetative yield of maize (Zea mays L.). Twosingle cross Corn Belt Dent maize hybrids were grown from sowingin a diurnal temperature regime of 16/6 °C day/night andin constant temperature environments of 16, 20, 24 and 28 °C.The 16/6 °C environment was close to the minimum for sustainedgrowth and 28 °C was near the optimum. Entire plants wereharvested at stages with 4, 6, 7 and 8 mature leaves in alltemperature treatments except 20 °C in which the final twoharvests were carried out at 9 and 10 mature leaves. Mean totalleaf number varied between 19.5 and 16.0 with the maximum occurringat 16/6 °C. Although harvests were carried out at comparableleaf numbers, and hence at similar developmental stages, thetime interval between sowing and harvest decreased considerablyas temperatures increased. The relative rates of dry weight and leaf area accumulationwith time increased with a Q₁₀ of 2.4 between 16 and 28 °C,while leaf appearance rate increased with a Q₁₀ of 2.9 overthe same range; both rates were highest at 28 °C. Althoughdry matter partitioning to the shoots increased with temperature,the area of individual leaves varied in a systematic patternwhich resulted in maximum leaf area, leaf area duration andconsequently dry weight being realized at 20 °C for anygiven stage of development. Zea mays, corn, low temperature stress, temperature response, growth, development     

The Effect of Temperature on Development and Dry-matter Accumulation of Vicia faba Seeds

Vicia faba plants were grown at 16 °C and the temperatureraised to 21 or to 26 °C, 51 d post anthesis (p.a.). Temperatureincrease accelerated pod and seed development, stimulated dry-matteraccumulation, starch and protein synthesis. However, the durationof dry-matter accumulation of seeds was shortened, resultingin a decrease of final seed weight. The effect of temperature on storage capacity (determined bycell number and cell size) was also investigated. Formationof new cells in the layer under the epidermis continued fora much longer period than previously has been described in theliterature. At all stages of development cotyledons containedyoung and small cells with small nuclei at the outside and theybecame larger and older towards the centre of the cotyledon.Cells at the centre were only able to divide at an early stageof seed development up to day 59 p.a. Thereafter cell divisionoccurred mainly in the first cell layer under the epidermisup to 63–67 d p.a. Nuclear counts of macerated cotyledoncells did not show an effect of temperature on the number ofcells. It has, however, been questioned if this method was suitableto measure accurately the number of cells at the last stagesof seed development. The rate of cell expansion was stimulated by higher temperaturesbut the duration of expansion was shortened and the final sizeof cells was not affected by the different temperature treatments.Senescence of the pod wall was accelerated at higher temperaturesand it is possible that transport of sucrose from the pod wallto the seeds terminated earlier than at lower temperatures.This may have resulted in an inhibition of cell formation atthe periphery of the cotyledons and in an inhibition of starchand protein synthesis. Vicia faba, protein, starch, cell size, cell number, temperature     

Day/night temperature environment affects cell elongation but not division in Lilium longiflorum Thunb

Lilium tongiflorum Thunb. cv. ‘Nellie White’ plantswere grown in different day/night temperature (DT/NT) environmentsto determine the anatomical basis for differential responsesof stem elongation to DT and NT. Lilium plants were forced in1986 and 1987 under 25 and 12 different DT/NT environments,respectively, with temperatures ranging from 14 to 30 °C.Parenchyma and epidermal cell length and width were measuredin stem tissue (1987) and epidermal cell length and width weremeasured in leaf tissue (1986). Total cell number per internodeand vertical cell number per internode were calculated. Stemparenchyma and stem and leaf epidermal cell length increasedlinearly as the difference (DIF) between DT and NT increased(DIF = DT —- NT), i.e. as DT increased relative to NT.DIF had no effect on stem parenchyma width, stem and leaf epidermalcell width, or cell number per internode. Data suggested thatstem elongation responses to DIF are elicited primarily througheffects on cell elongation and not division. Key words: Thermoperiodism, thermomorphogenesis, stem elongation, DIF, cell division, cell elongation, leaf expansion     

Difference in light-induced increase in ploidy level and cell size between adaxial and abaxial epidermal pavement cells of Phaseolus vulgaris primary leaves

Changes in nuclear DNA content and cell size of adaxial andabaxial epidermal pavement cells were investigated using brightlight-induced leaf expansion of Phaseolus vulgaris plants. Inprimary leaves of bean plants grown under high (sunlight) ormoderate (ML; photon flux density, 163 µmol m^–2s^–1) light, most adaxial epidermal pavement cells hada nucleus with the 4C amount of DNA, whereas most abaxial pavementcells had a 2C nucleus. In contrast, plants grown under lowintensity white light (LL; 15 µmol m^–2 s^–1)for 13 d, when cell proliferation of epidermal pavement cellshad already finished, had a 2C nuclear DNA content in most adaxialpavement cells. When these LL-grown plants were transferredto ML, the increase in irradiance raised the frequency of 4Cnuclei in adaxial but not in abaxial pavement cells within 4d. On the other hand, the size of abaxial pavement cells increasedby 53% within 4 d of transfer to ML and remained unchanged thereafter,whereas adaxial pavement cells continuously enlarged for 12d. This suggests that the increase in adaxial cell size after4 d is supported by the nuclear DNA doubling. The differentresponses between adaxial and abaxial epidermal cells were notinduced by the different light intensity at both surfaces. Itwas shown that adaxial epidermal cells have a different propertythan abaxial ones. Key words: Cell enlargement, endopolyploidization, epidermal pavement cells, incident light intensity, leaf expansion, nuclear DNA content, Phaseolus vulgaris     

Grass Leaf Elongation Rate as a Function of Developmental Stage and Temperature: Morphological Analysis and Modelling

Elongation of successive leaves was measured following defoliationof tall fescue plants in controlled environments. Measurementswere made under constant temperatures of 24 °C and 14 °C,and after temperature changes from 24 to 14 °C andvice versa.A morphological analysis of the growing leaf was made from thetime it was 1 mm long until it was fully elongated. The timeelapsed from initiation until the leaf was 1 mm long was estimated.Young leaves less than 1.5 mm long elongated slowly at a constantleaf elongation rate (LER). By extrapolating this LER back toleaf initiation from the apex it was calculated that elongationlasted 42.5 d at 24 °C and 51 d at 14 °C. Lengths ofthe division zone (DZ) and the extension-only zone (E-OZ) increasedto a maximum and then decreased during leaf development. Temperaturechange had an immediate effect on LER but the response varieddepending on the direction of the temperature change. To describethese different features, an empirical model of DZ and E-OZwas designed. Its five parameters were optimized at constanttemperature. The model was then used to simulate the LER ofplants subjected to temperature changes. Instant and lastingeffects of the initial temperature on mean LER in plants transferredfrom 14 to 24 °C andvice versawere well simulated. It wasconcluded that the major reason for differences was due to thegrowth stage (DZ and E-OZ lengths) at which the changes occurredat both temperatures.Copyright 1999 Annals of Botany Company Festuca arundinaceaSchreb., tall fescue, growth zone, division zone.     

Cell Division and Expansion in the Growth of the Leaf

Volumes and numbers of cells were determined at different stagesof development of the fifth leaf of Lupinus albus, and eachof the second pair and the tenth leaf of Helianthus annuus.In the case of the second pair of sunflower leaves the valuescover the whole life of the leaf from initiation to senescence. During both primordial development and the ensuing ‘grandperiod of growth’ division is the determinant of growth.About 10 per cent. of the cells in the fully grown leaf arelaid down before leaf-emergence; the remaining 90 per cent.are formed during unfolding. Division does not cease in thelupin leaf or the second pair of sunflower leaves until theyhave reached half their maximum area. The tenth leaf, on theother hand, is as much as three-quarters fully grown beforedivision ceases. Cell expansion commences soon after leaf initiation and continuesthroughout the life of the leaf. With lupin and the second pairof sunflower leaves there is a fourfold increase in the averagevolume of the cells before emergence from the apical region.During unfolding, there is a further tenfold increase in theaverage volume of the cells of the lupin leaf, and a twentyfoldincrease with the second pair of sunflower leaves. Expansioncontinues after the cessation of division but this further increasein volume is comparatively small. The data are discussed in relation to the ‘two phase’hypothesis of leaf development.     

Developmental Physiology of Sugar Beet: I. THE INFLUENCE OF LIGHT AND TEMPERATURE ON GROWTH

Sugar beet plants were grown for 12 weeks from emergence ingrowth rooms at temperatures of 10, 17, 24 and 31 °C and20, 50, 80, and 110 cal visible radiation cm^-2d^-1, and the changeswith time in their dry weight, leaf area, leaf numbers, andstorage root sugar determined. The first stage of growth wasdominated by the development of the shoot, but the storage rootgradually assumed increasing importance and eventually grewat a faster rate and to a greater weight than the shoot. Therelative growth rate and final yield of dry matter of the shootwere greatest at 24 °C and of the root between 17 and 24°C. The relative rate of expansion and the final area ofthe leaf surface were also greatest at 24 °C, whilst therates of production and of unfolding of leaves were greatestat about 17 °C. All these attributes were increased withincreased radiation. Net assimilation rate increased almostproportionately with radiation and was not significantly affectedby temperature.The relationships of total leaf area with plantdry weight, root dry weight with shoot dry weight, and totalleaf number with plant dry weight were scarcely affected bychanges in radiation, but were much influenced by temperature.Plants of the same dry weight generally had bigger roots andsmaller areas of leaf surface as temperatures departed from24 °C and had most leaves at 17 °C. Sugar concentrationsin the storage root were greatest at 17 °C, but the totalamount of sugar was about the same at 17 and 24 °C. Theconcentration of sugar in the storage root depended on rootsize.Thus, temperature affected both the rate and pattern ofdevelopment, and radiation affected the rate but not the patternof development.     

Leaf Growth in Phaseolus vulgaris: I. Growth of the first pair of leaves under constant conditions

The growth of the first pair of leaves of Phaseolus vulgaris(French bean) has been studied during germination and followingemergence of the seedling. The leaves are well developed inthe embryo and, at 22.5° C, show an exponential increasein fresh weight, dry weight, and leaf area up until about eightdays from planting. Cell division commences about two days afterplanting and is exponential for a short period. Considerablechanges in cell volume occur during the period over which celldivision occurs. Cell division ceases soon after emergence andunfolding, when the leaf has reached only 17 per cent of itsfinal area. Cessation of cell division is followed by a phaseof growth which is due entirely to cell expansion. The significanceof these findings is discussed in relation to recent work onother genera.     

Expression of Vernalization Genes in Near-isogenic Wheat Lines: Effects of Night Temperature

Four near-isogenic lines of wheat (Triticum aestivum L.em Thell)were used to compare selected night temperatures for their effectivenessas vernalizing temperatures. All treatments (conducted withina phytotron) had a common day temperature of 20 °C for 12h and night temperatures were 4, 7, 10, 13 and 20 °C. Interpretationof results for reproductive development was confounded by threeinteracting factors, their relative importance varying withgenotype. Firstly, development rate was generally slower atlower night temperatures. Secondly, in contrast, there was atendency for lower night temperatures to hasten developmentrate if vernalization requirements were satisfied. Thirdly,the lower night temperatures provided a more favourable environmentfor leaf production such that for some genotypes, vernalizedplants had higher final leaf numbers than unvernalized plants.Only for the genotype with the strongest vernalization response(vrn1 vrn2) did hastening of development due to vernalizationoverride any delaying effects. For this genotype, 4, 7 and 10°C were vernalizing temperatures. For the other three genotypes,any hastening of development due to vernalization was outweighedby delaying effects of lower night temperatures. Spikelet numberand days to anthesis were positively correlated in three ofthe four genotypes. It appeared that differences in spikeletnumber were a direct result of night temperature influencingthe duration of the spikelet phase and/or rate of spikelet initiation.Plant size at flowering was determined by the differential effectsof night temperature on growth and development rates. Triticum aestivum L., wheat, vernalization, night temperature, isogenic lines     

The Roles of Cell Division and Cell Expansion in the Growth of Alfalfa Leaf Mesophyll

Leaf mesophyll of Medicago sativa (L.) was investigated to determinethe roles of cell division and cell expansion in tissue growth.Samples of leaf tissue were macerated, stained, and squashed.The slides were studied under a phase microscope to determinethe percentage of recently divided cells and the average celldiameter for leaflets of varying lengths. Cell division wasgreatest in young leaflets and virtually ceased as a leaf lengthof 12 mm was attained. For leaflets less than 12 mm in length,the rate of increase in cell size appeared to be inversely associatedto the degree of cell division. For alfalfa leaflets greaterthan 12 mm in length, the mean cell size increased in proportionto leaf length since cell division had virtually ceased.     

Cell growth in expanding primary leaves of Phaseolus

Plants were grown at 25 and 20° C in 6, 12, and 18 h daylengths.Final area of the primary leaf pair ranged from 105 to 209 cm²,and for a given temperature was greatest in the 12 h and leastin 6 h daylength. Cell numbers per leaf were similar for alltreatments. In the 6 h daylength leaves were thinner, containedless chlorophyll and ethanol-insoluble dry matter, and had considerablysmaller cells than leaves on plants in the longer daylengths;final levels of protein and cell-wall material per cell werealso low, although levels of nucleic acid per cell were as highas, or higher than, those for leaves in 12 and 18 h days. Itis concluded that the low levels of protein and cell-wall materialare associated with a low level of photosynthesis, and thatthe small area of these leaves is a result of the reductionin cell size. In the 12- and 18-h daylenghts, protein and cell wall per cellincreased linearly with time, and when expansion of the laminawas completed, values for these parameters were found to besimilar. Cell size, as measured by fresh weight, was also similarat this stage, although small differences in lamina thicknesswere found. Thus the smaller area for leaves in 18-h days wasnot due to a reduction in mean cell size, although differencesin epidermal cell dimensions must be involved. From consideration of simple models it is concluded that increasein cell wall material during lamina expansion is associatedwith increase in wall area, but that the continued formationof wall material after lamina expansion has ceased is accountedfor by deposition on already existing walls. This continuedincrease in wall material occurs at a time when protein andnucleic acid levels per cell are declining.     

Low Temperature Affects Pattern of Leaf Growth and Structure of Cell Walls in Winter Oilseed Rape (Brassica napus L., var. oleifera L.)

Three-week acclimation of winter oilseed rape (Brassica napusL. var. oleifera L.) plants in the cold (2 °C) resultedin a modified pattern of leaf cell enlargement, indicated bythe increased thickness of young leaf blades and modified dimensionsof mesophyll cells, as compared with non-acclimated tissuesgrown at 20/15 °C (day/night). The thickness of leaf cellwalls also increased markedly during cold acclimation but itdecreased in response to a transient freezing event (5 °Cfor 18 h followed by 6 or 24 h at 2 °C, in the dark). Cellwalls of the upper (adaxial) epidermis were most affected. Theirultrastructure was modified by cold and freezing treatmentsin different ways, as revealed by electron microscopy. Possiblereasons for the cold- and freezing-induced modifications inthe leaf and cell wall morphology and their role in plant acclimationto low temperature conditions are discussed. Copyright 1999Annals of Botany Company Acclimation, Brassica napus var. oleifera, cell wall ultrastructure, cold, freezing, leaf structure, winter oilseed rape.     

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     

Studies on the Expansion of the Leaf Surface: III. THE INFLUENCE OF RADIATION ON CELL DIVISION AND LEAF EXPANSION

The numbers of leaves and the areas and cell numbers of leavesfrom the first five nodes of the cucumber were determined throughouttheir development with three different amounts of daily radiationand two conditions of nutrient supply. The rate of leaf production was found to be constant with timefor any one amount of radiation and to increase with increasedradiation. The transition from the rate in darkness to thatin light was sharp and occurred more quickly the higher theradiation. Provided the nutrient supply was high the ultimate areas ofindividual leaves were greater the higher the radiation; ifmineral nutrients were depleted the maximum area of a leaf occurredwith an intermediate amount of light. This arose because thesefactors exercised a differential effect on the various phasesof growth. Each leaf commenced its life as a mass of dividing cells andthe mean rate of division remained constant until it unfoldedfrom the terminal bud. The mean rate of division was much greaterat high than at low levels of radiation and was interpretedas being regulated by carbohydrate supply. Although expansionof some cells was likely before unfolding, after this stagethere was a marked decrease in the proportion of cells proceedingto division. The duration of division in the leaf after unfoldingwas independent of radiation; although 70–98 per cent,of the final number of cells were formed after unfolding, theultimate number was effectively determined by the rate of divisionprior to unfolding. Low nutrient supply restricted the duration of division in theleaf to a significant extent and the expansion of cells to avery considerable degree. The smaller leaves on plants receivinghigh rather than medium radiation were due to their cells beingmuch smaller.