Studies on the Expansion of the Leaf Surface: V. CELL DIVISION AND EXPANSION IN A DEVELOPING LEAF AS INFLUENCED BY LIGHT AND UPPER LEAVES

The development of the third leaf was followed from soon afterinitiation to the attainment of full area in cucumbers grownunder two levels of radiation and, following leaf unfolding,with the shoot above the developing leaf present or absent.Cell numbers, proportion of cells in meta-, ana-, and telo-phases,leaf areas, and aspects of leaf structures were recorded. Cell numbers increased exponentially until near unfolding atboth intensities—at a higher rate under high light andfor a longer time under low light. Thereafter, the rate declinedto zero. Final cell numbers were higher under high than lowlight. Leaf structure and cell dimensions parallel to the leafsurface were not affected by light intensity and therefore finalareas depended solely on cell numbers. Apex removal resultedin much larger cells, and in consequence, correspondingly greaterleaf areas. The proportions of cells in the mitotic stages recordedwere similar at both intensities but fell steadily throughoutthe periods of exponential increase of cell number. Previous hypotheses that all cells are active in division untilthe leaf unfolds could be largely reconciled with mitotic behaviourunder high light, but some decline in the proportion of dividingcells was indicated. Under low light, such a decline must beimportant. Greater cell numbers in leaves at high light arose,therefore, from a higher rate of division in dividing cells,combined with a larger proportion remaining active. Greater leaf areas following apex removal are consistent withhypotheses of competition between developing leaves but throwno new light on the specific factors involved.     

Studies on the Expansion of the Leaf Surface: I. THE INFLUENCE OF TEMPERATURE

An investigation was made of the expansion of the leaf surfaceof cucumber at temperatures of 12°, 18°, 24°, and30°C. with two levels of visible radiation (40 and 80 cal.cm.^–2 day^–1). The relative rate of expansion ofthe leaf surface increased with temperature up to 24° butwas lower at 30° than at 24°. It was slightly greaterwith the higher than the lower level of radiation at the lowertemperatures only. This rate was the resultant of the rate ofunfolding of new leaves and the rate of expansion of the componentleaves. The rate of leaf production increased with increasingtemperature up to 24° and was constant there-after, butleaves unfolded from the terminal bud more rapidly with increasein temperature over the entire range. The rate of expansionof individual leaves was greatest at 24°, being less atboth lower and higher temperatures. Differences in this ratebetween temperatures increased in the order: cotyledon, leaf1, leaf 2. Leaf production and unfolding was greater with thehigher level of radiation but the expansion of individual leaveswas not influenced. These results suggested the following interpretation of theexpansion the leaf surface. Its potential rate is set by therate of unfolding of leaves from the terminal bud, which dependsmainly on the temperature and the rate of assimilation by theupper leaves and the terminal bud, the demand for assimilateexceeding the supply in this region. The demand for mineralsubstrates by the terminal bud is low and not influenced bya wide variation in potential supply. After unfolding from theterminal bud, the leaf provides most of its own supply of carbohydrateby assimilation and this can be met at a low level of radiation.Surplus assimilate is diverted to the roots and stems whichrespond much more to increased radiation than does the leafsurface. The demand for mineral substrates by expanding leaves,however, is high—the greater the number expanding at anytime the more likely is the demand by any one leaf to exceedthe supply. This leads to a reduction in the number of celldivision and, consequently, a reduced rate of expansion anda smaller leaf. The optimum level of any environmental factoris that at which the most effective compromise between theseconflicting processes is reached.     

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.     

LEAF DEVELOPMENT OF RUMEX PATIENTIA L. (POLYGONACEAE) EXPOSED TO UV IRRADIATION (280-320 NM)

Two factors which affect leaf ontogeny and ultimate leaf size: (1) the rate and duration of cell expansion, and (2) the rate and duration of cell division, were examined for their role in the slower early leaf growth rate and the smaller size of fully expanded leaves of plants exposed to ultraviolet-it (UV-B 280-320 nm) radiation. Rumex patientia L. was grown in controlled environment chambers under enhanced UV-B radiation (equivalent to daily solar UV-B irradiation at 40°N latitude in mid-May with an atmospheric ozone concentration of 0.20 atm-cm) and control treatments. The pattern of growth as expressed in changes of mean cell size in two distinct cell types, tissue cell density, and length of the entire leaf blades are consistent with the hypothesis that the radiation primarily affects cell division rather than cell expansion. Furthermore, it appears that the radiation probably alters the rate rather than the duration of the cell division phase. An understanding of the mechanism of radiation damage should facilitate prediction of how this stress may interact with other stresses to which plants are normally subjected. Species with normally prolonged periods of cell division during leaf expansion may be particularly impacted if solar UV radiation were intensified as a result of atmospheric ozone reduction.     

Ultraviolet-B radiation reduces the rates of cell division and elongation in the primary leaf of wheat (Triticum aestivum L. cv Maris Huntsman)

The primary leaf of wheat (Triticum aestivum L. cv Maris Huntsman) was used as a model system to examine how elevated ultraviolet‐B (UV‐B; λ= 280–320 nm) radiation affected growth. A reduction in the rate and duration of growth of the primary leaf, in response to UV‐B, was the result of changes in both the rate and extent of cell division and elongation. UV‐B reduced the proportion of mitotically active cells (mitotic index) and increased the time taken for cell division (cell doubling time). Thus the supply of cells into the elongation zone was reduced, and this, coupled to a reduction in the rate of elongation, resulted in reduced leaf growth. This analysis of the spatial distribution of growth provided a means of calculating the age of cells within the leaves. Cells of UV‐B‐treated leaves were found to age more quickly than those of the controls. This analysis will enable future studies to take account of age‐related changes when interpreting the response of plants to any number of environmental stresses that affect leaf development.     

Nitrate availability and shoot area development in small willow (Salix viminalis)

Abstract. The effect of nitrate supply upon leaf area development in willow ( Salix viminalis ) was investigated. Rooted cuttings were grown in culture solution at two different rates of exponentially-increasing nitrate supply and with free access to other nutrients. During the lag-phase in attaining stable nutrition, the rate of production of unfolded leaves, in the treatment with poorer N-supply, decreased. After acclimation to the different N-availabilities, the rate of dry matter increase per plant nitrogen was constant and the same in both treatments. The ratio of total leaf area to total amount of nitrogen in the plant was slightly higher in the treatment with poorer nitrogen supply. During the exponential (stable) phases of growth, nearly all the increase in total leaf area was attributable to main-stem leaves; large numbers of leaves on axillary shoots in the treatment with better N-supply contributed little to the total area. During the stable growth phase, the rate of production of unfolded leaves was the same in both treatments. Higher maximum values of relative rate of increase in area of single leaves were found with a better N-supply and at higher leaf positions on the main stem. The duration of leaf expansion was not much affected by N-supply. Final leaf size was greater at higher positions on the main-stem. Within a treatment (investigated for poorer N-supply), differences in final leaf size were proportional to differences in numbers of epidermal cells. Final leaf size was greater, at the same node, in the treatment with better N-supply. Better N-availability had increased the final size but not the number of epidermal cells. It was concluded that the higher exponential rate of total area increase with a better N-supply was largely associated with higher rates of expansion in epidermal cells.     

The Effect of Temperature on the Growth of Individual Leaves of Vicia faba L. in the Field

The generalized logistic curve was used to describe the growthof individual leaves in crops of Vicia faba L. Durations of.expansionand mean absolute growth rates were derived from these curves.The duration of expansion was inversely related to temperatureaveraged over four days from unfolding. This relationship wasindependent of leaf position except for the lowest leaves. Theduration of expansion of a leaf was related to the rate of productionof new leaves, the number of expanding leaves remaining relativelyconstant. Absolute growth rates varied with leaf position upto leaf 10. At higher leaves, in the absence of water stress,absolute growth rate was a function of temperature and radiation. Vicia faba L., field bean, leaf growth, temperature     

Devlopmental Physiology of Sugar-Beet: II. EFFECTS OF TEMPERATURE AND NITROGEN SUPPLY ON THE GROWTH, SOLUBLE CARBOHYDRATE CONTENT AND NITROGEN CONTENT OF LEAVES AND ROOTS

Plants were grown at temperatures of 15 and 25 ?C with two ratesof nitrogen supply. The changes in dry weight, leaf area, cellnumber, mean cell volume, soluble carbohydrate, and total nitrogenconcentration of the cotyledons, the first and second pair oftrue leaves, and the storage root were measured. Changes incell number and cell volume of the first pair of true leavesand storage root of plants were also measured at 11, 18, 25,and 32 ?C. Leaf growth before unfolding was chiefly by increase in cellnumber and after unfolding by increase in mean cell volume,while the growth of the storage root was almost entirely byincrease in cell number. The rates of cell division and cellexpansion were fastest at 25 ?C, but the initially high ratesof cell division in the terminal bud and in individual leavesdecreased rapidly and greater rates were maintained at the sub-optimaltemperatures, i.e. 15 and 18 ?C. After an initial period ofslow growth, the first-formed leaves grew faster and becamelarger at 15 than at 25 ?C. Leaves were produced, unfolded,grew faster, and became larger with increase in the externalconcentration of nitrogen, because cells divided and expandedfaster, so that nitrogen increased the number and size of cells. Sugar concentration was greater at 15 than at 25 ?C in leavesbut not in the storage root. Sugar concentration in the petiolesof the first and second pair of true leaves increased to 1.2and 2.0 per cent fresh weight respectively. Decreased nitrogensupply temporarily increased the sugar concentration of cotyledonpetioles and the seedling hypocotyl, but later decreased itin the leaves and storage root. Nitrogen concentration was greaterin the leaves and storage root at 15 than at 25 ?C with thelarger nitrogen supply. Nitrogen concentrations were similarin young leaves of all treatments but as the size of leavesincreased nitrogen concentrations decreased most rapidly at25 ?C with the smaller nitrogen supply. It is suggested that when increased leaf production and storage-rootgrowth occurs at temperatures below the growth optimum (25 ?C),they may be due to an effect of increased carbohydrate supplyon cell division and sugar storage.     

Leaf dynamics,self-shading and carbon gain in seedlings of a tropical pioneer tree

We examined leaf dynamics and leaf age gradients of photosynthetic capacity and nitrogen concentration in seedlings of the tropical pioneer tree, Heliocarpus appendiculatus, grown in a factorial design under controlled conditions with two levels each of nutrients, ambient light (light levels incident above the canopy), and self-shading (the gradient of light levels from upper to lower leaves on the shoot). Correlations among these parameters were examined in order to determine the influence of self-shading, and the regulation of standing leaf numbers, on leaf longevity and its association with leaf photosynthetic capacity. Leaf longevity and the number of leaves on the main shoot were both reduced in high light, while in the low light environment, they were reduced in the steeper self-shading gradient. In high nutrients, leaf longevity was reduced whereas leaf number increased. Leaf initiation rates were higher in the high nutrient treatment but were not influenced by either light treatment. Maximum-light saturated photosynthetic rate, on an area basis, was greater in the high light and nutrient treatments, while the decline in photosynthetic capacity in realtion to leaf position on the shoot was more rapid in high light and in low nutrients. Leaf longevity was negatively correlated among treatments with initial photosynthetic capacity. The leaf position at which photosynthetic capacity was predicted to reach zero was positively correlated with the number of leaves on the shoot, supporting the hypothesis that leaf numbers are regulated by patterns of self-shading. The negative association of longevity and initial photosynthetic capacity apparently arises from different associations among gradients of photosynthetic capacity, leaf numbers and leaf initiation rates in relation to light and nutrient availability. The simultaneous consideration of age and position of leaves illuminates the role of self-shading as an important factor influencing leaf senescence and canopy structure and dynamics.     

Endogenous Short Period Rhythms in the Movements of Unifoliate Leaves of Phaseolus angularis Wight

Rhythmic rotational movements with the midvein as the axis have been observed in the unifoliate leaves of Phaseolus angularis Wight grown under controlled environmental conditions with continuous light. The mean period of this movement for all leaves was 53.2 ± 4.3 minutes and remained constant as the leaf matured, except after removal of the apical meristem and emerging trifoliate leaf when the period increased by about 5 minutes. The amplitude of the movement also remained constant as the leaf matured. These rotational movements were pronounced when the leaf blade was in a horizontal position and were not evident during the downward or “sleep” movements of the leaf. This movement began 3 days after leaf unfolding and continued for at least 6 days. It was most pronounced at the time of inflection of the leaf length growth curve after the logarithmic phase of growth.     

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     

温室甜椒叶面积指数形成模拟模型

叶面积指数是光合作用驱动的作物生长模型以及冠层蒸腾模型所需的重要作物参数,温度和辐射是影响叶片生长的重要环境因子.通过不同定植期、不同品种、不同地点的试 验,定量分析了温室甜椒出叶数、叶片长度和叶面积指数与温度和辐射的关系,构建了温室 甜椒叶面积模型,并利用独立的试验资料对模型进行了检验.结果表明：甜椒出叶数与出苗 后累积辐热积呈指数函数关系;叶片长度与出叶后累积辐热积呈负指数函数关系;甜椒出叶 数、叶片长度和叶面积指数的模拟结果与实测值之间的决定系数R²分别为0.94、0.89、0.93,其回归估计标准误RMSE分别为3.4、2.15 cm、0.15.该模型能够利用气温、辐射、 种植密度和出苗日期准确地预测温室甜椒叶面积指数动态,且模型参数少、实用性强,可以为温室甜椒生长模型和蒸腾模型提供必需的叶面积指数动态信息.     

Recovery of leaf area through accelerated shoot ontogeny in thrips-damaged cotton seedlings

BACKGROUND AND AIMS: Leaf area of cotton seedlings (Gossypium hirsutum) can be reduced by as much as 50 % by early season thrips infestations, but it is well documented that plants can regain the difference in leaf area once infestation ceases. The processes involved in the recovery have not been identified. Hypotheses include enhancement of the photosynthetic rate of the damaged leaves, more efficient leaf construction (i.e. more leaf area per unit of dry matter invested in new leaves), and more branching. METHODS: This 2-year field study examined these hypotheses and found that thrips-affected plants recovered from a 30 % reduction in total leaf area. During the recovery period, repeated measurements of gas exchange, leaf morphology and individual leaf areas at all nodes were made to assess their contribution to the recovery. KEY RESULTS: Recovery was not achieved through the previously proposed mechanisms. The pattern of nodal development indicated that the duration of leaf expansion of the smaller deformed leaves was shorter than that of control leaves, possibly because they had fewer cells. The production and expansion of healthy upper node leaves in thrips-affected plants could, therefore, begin sooner, about 1-2.5 nodes in advance of control plants. The proposed process of recovery was evident but weaker in the second year where thrips numbers were higher. CONCLUSIONS: It is concluded that thrips-affected plants overcame the leaf area disparity through an accelerated ontogeny of main stem leaves. By completing the expansion of smaller but normally functioning lower node leaves earlier, resources were made available to the unfolding of larger upper node leaves in advance of control plants. The generality of this mode of plant resistance in pest damage remains to be determined.     

Persistent effects of seedling treatment on growth of sugar beet in pots

Further evidence is provided that the environment of sugar-beet seedlings, or growth substances applied to seedlings, continues to influence growth when the plants are later in other environments. Sugar-beet seeds were germinated at 20 °C in 8, 16 or 24 h photoperiods of constant light intensity, i.e. with different amounts of total radiation. When the seedlings had two leaves (15–18 days old) they were transferred to large pots in the glasshouse. Some seedlings were treated with (2-chloroethyltrimethylammonium chloride) either sprayed on the leaves or applied to the soil, at different times. The treatments affected areas of individual leaves throughout the growing period; plants raised in 24 h photoperiod had the largest leaves, and those in 8 h photoperiod the smallest. Consequently, 24 h plants had most dry matter and 8 h plants least. Plants given most radiation produced leaves fastest and CCC applied early increased the rate, but as the leaves were smaller, except late in 1967, and died sooner, the leaf area duration was less and so yields were less. CCC applied later did not affect leaf production. There was no interaction between amount of radiation and CCC treatment. Twenty-four hour plants had the greatest net assimilation rate (E) early. CCC decreased E early, but increased it later and more when sprayed on the leaves than when applied to the soil. Some factor, possibly pot size, eventually restricted growth and probably diminished the effect of the treatments applied to the seedlings.     

Changes in chloroplast number per cell during leaf development in spinach

Summary The amounts of chlorophyll and nitrogen and the numbers of cells per unit area change as the green leaves of spinach plants grow and increase in size in the light. The changes in the numbers of chloroplasts per cell were measured by a new method. A 5-fold increase in the numbers of chloroplasts per cell took place in both palisade and mesophyll cells over a growing period of 10 days during which time the area of the leaves increased from 1 to 50 cm². Proplastids were not present in the young green leaves but electron-microscope and phase-contrast observations showed the presence of grana-containing chloroplasts, many of which appeared to be undergoing division by constriction. It is suggested that the large increase in chloroplast numbers as leaf cells grow and expand in the light is from the division of differentiated chloroplasts containing grana.     

Leaf growth of white mustard (Sinapis alba) in different environments

Summary The numbers of cells and area of fully expanded leaves were determined on successive leaves of Sinapis alba grown either in 8 hr. photoperiod (vegetative plants) or 16 hr. photoperiod (flowering plants) at a constant temperature of 20°C. In the 8 hr. photoperiod leaf 9 had the greatest area but leaf 12 had most cells. In 16 hr. photperiod leaf 5 had the greatest area but leaf 9 had most cells. The relationship between area and cell number of successive leaves on the main stem fell into 3 distinct phases: in phase (1), cell number increased at a greater rate than leaf area; in phase (2), leaf area decreased while cell number increased; in phase (3), cell number and leaf area decreased proportionally. For an increase in unit area, cell number increased more in 8 hr. than in 16 hr. photoperiod.Using final area and final cell number of successive leaves, by extrapolation the cell number of unit area of primordium has been deduced. Cell number per unit area increased in successive primordia up to a certain node after which it remained constant at succeeding nodes. It was found that in plants grown under different conditions the cell number per unit area in successive primordia increased at a constant logarithmic rate. That is, cells became progressively smaller. It is concluded that changes in cell size of successive primordia are not influenced by the environment but are under internal control.     

Bean Leaf Expansion in Relation to Temperature

When dwarf Phaseolus vulgaris plants were grown in a controlledenvironment at 20, 25, 30, and 35° C, expansion of the primaryleaves occurred in two phases with an intermediate lag. Varyingrates and duration of expansion were involved, leading to greatestfinal areas at the two intermediate temperatures. Dry weightsof the leaves and leaf areas were similary influenced by temperature,except that the initial rates of increase continued for a longerperiod for weights than for areas. The rates of cell divisionand final numbers of cells were similar from 25 to 35° C,but both were decreased at 20° C. Final cell sizes were,on the other hand, decreased only at the highest temperature.The time trends of cell expansion varied greatly with temperature. Leaf expansion is discussed as a possible consequence of substratesupply, which may be determined by temperature in a number ofways. Cell division and cell expansion are not considered tobe joint direct determinants of leaf expansion. Temperatureinfluences division, with two consequences; the rate interactswith substrate supply to determine size of cells, and finalcell number affects potential leaf area. Cell size is regardedas being secondary to numbers of cells and total material available,although some factors can vary cell size independently of substrate,e.g. water status. An important control of leaf growth, until the attainment ofabout half the final area, may be exercised by way of the leaf.Subsequently, intra-plant competition is likely to dominate.     

Studies on the Expansion of the Leaf Surface: II. THE INFLUENCE OF LIGHT INTENSITY AND DAYLENGTH

The parts played by constant amounts of visible radiation perday and its two components—daylength and intensity—ininfluencing the growth of Cucumis sativus have been investigated.The amount of radiation per day had a far greater influencethan either of its components per se. Nevertheless, small significanteffects of photoperiod were found, leaf expansion and dry weightincrease being greatest at daylengths between 10 and 15 hr.rather than with longer days which, with similar daily totals,would be expected to give the greatest amounts of assimilation. Rates of leaf production and appearance were greatest with thehighest amounts of radiation, but the rates of expansion ofindividual leaves and their maximum areas were greatest withintermediate amounts of radiation. This response resulted inan optimum curve relating the leaf surface and the dry weightattained after a given period to radiation. The amount of radiationgiving the maximum leaf surface and dry weight decreased withage and with external nutrient supply, but at any one age washigher for increase in dry weight than for increase in leafsurface; stem and root tissues responded more to high radiationthan did the leaf surface. The net assimilation rate was a linearfunction of visible radiation over the range of 15–120cal. cm.^-2 day^-1 explored, the highest value of radiation usedrepresenting the intensity at which photosynthesis would beexpected to be maximal over a 12–15 hr. day. The inhibitory effect of high radiation on leaf expansion andthe resultant influence on the growth of the plant are explainedin terms of the number and intensity of ‘sinks’for carbohydrate and mineral nutrients within the plant.     

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 modular character of growth in Nicotiana tabacum plants under steady-state nutrition

The impact of different plant growth rates on biomass allocation and growth distribution in tobacco was studied on the whole plant, total leaf area and single leaf level. On the whole plant level, constant relationships were found between the total leaf area and the biomass allocation to leaves and the nonphotosynthetic organs (roots and stem) independent from the overall growth rate and the nutrient addition rate to the plants. On the level of total leaf area, plants grown at lower nutrient supply reached a distinct distribution of leaf area later than those grown at higher nutrient supply, but the normalized distribution of leaf area along the stem at a certain plant size did not differ between plants growing at different nutrient supply and growth rates. On the leaf blade level, growth rates declined, initially linearly, from the leaf base to the leaf tip. Distinct gradients within the side veins were not observed, but the growth rates of the side veins were closely correlated to the adjacent mid-vein segments. These gradients flattened with increasing size of the leaf. The modular character of growth in tobacco is discussed in the context of basic growth analysis and as a framework for physiological, cytological, biochemical, and molecular studies in growing plants.Key words: Nicotiana tabacum, whole plant, total leaf area, leaf growth, growth rate, biomass.