ACROPETAL LEAF DIFFERENTIATION IN QUERCUS RUBRA (FAGACEAE)

Northern red oak (Quercus rubra L.) leaves were shown to mature progressively from base to tip of the lamina based on studies of growth rates, anatomical differentiation, and ¹⁴C-transport. Lamina expansion in both length and width ceased in the basal quarter of the leaf before the apical quarter. Cell expansion and tissue differentiation were more advanced at the base than at the tip of leaves at 10%–20% of full expansion. Physiological data supported the morphological and anatomical data. Sink activity was examined by following the distribution of ¹⁴C imported into sink leaves with direct vascular connections to the source leaf to assure uniform assimilate supply to the sink leaves. Leaves approximately 50% of full expansion imported five to seven times more ^l4C-assimilates into the tip than into the base of the leaf, consistent with continued sink activity in the leaf tip after import by the leaf base has ceased. Transport of ¹⁴C from portions of the leaf, indicating source activity, occurred first in the basal portion of the lamina. The base functioned as a source at approximately 40% of full expansion; the tip, at approximately 60%. Thus, northern red oak displays an acropetal pattern of leaf expansion and differentiation, unlike the more typical pattern of basipetal leaf development defined in many other dicotyledonous genera with simple leaves.     

Distribution of imported 14C in developing leaves of eastern cottonwood according to phyllotaxy

Summary Individual leaves of eastern cottonwood (Populus deltoides Bartr.), representing an ontogenetic series from leaf plastochron index (LPI) 3.0 to 8.0, were fed ¹⁴CO₂ and harvested after 2–24 h. Importing leaves from LPI-1.0 through 8.0 on each plant were sectioned into 9 parts, and each part was quantitatively assayed for ¹⁴C activity. The highest level of ¹⁴C import was by leaves from LPI 1.0 to 3.0, irrespective of source-leaf age. ¹⁴C was translocated preferentially to either the right or left lamina-half depending on the position of the importing leaf in the phyllotactic sequence and its stage of development. For example, import was high when the importing leaf and the source leaf had two vascular bundles in common, moderately high with one bundle in common, and low with no bundles in common. The distribution of ¹⁴C within young importing leaves was highest in the lamina tip and decreased toward the base. With increasing leaf age, incorporation declined in the lamina tip and increased in the base.It may be concluded that each cottonwood leaf progresses through a continuum of importing and exporting stages as its lamina expands. The photosynthate imported by a given leaf is compartmentalized, with different exporting leaves supplying photosynthate to rather restricted regions of the lamina. Such localization within the importing leaf depends on its vascular connections with each of the exporting leaves, and these are predictable from a knowledge of the phyllotaxy.Plant Physiologists.     

QUANTITATIVE ANALYSIS OF LEAF DEVELOPMENT IN XANTHIUM PENSYLVANICUM

Maksymowych , Roman . (Villanova U., Villanova, Pa.) Quantitative analysis of leaf development in Xanthium pensylvanicum. Amer. Jour. Bot. 46(9): 635–644. Illus. 1959.—An attempt was made to find a quantitative way of describing the development of the leaf and to correlate the developmental processes, designating precisely their sequence. The processes were presented in terms of the absolute and relative rates of leaf length, expansion of lamina in surface, increase in thickness, rates of cell division of leaf 9 and 13, and tissue differentiation of 3 portions of the lamina. All rates were estimated over the entire period of development, from initiation of a primordium to its maturity. The leaf plastochron index (L.P.I.) was used as a morphological time-scale. The relative plastochron rates were used for the purpose of correlation of the developmental processes. Leaf 9 elongates exponentially up to 3.0 L.P.I. with an average relative rate (dlnL/dpl) of about 0.78 pl^-1, and it stops growing around 8.0 L.P.I. The lamina stops elongating about 1.5 plastochrons before the petiole. The tip of the lamina expands its surface at a constantly lower relative rate than the middle and the basal portions of the blade. The average relative rate of expansion in area (dlnA/dpl) for the whole lamina is 1.7pl^-1 during the exponential stage. Differentiation of the laminar tissues proceeds basipetally, from the tip toward the base of the leaf. The relative rate of expansion of lamina in thickness (dlnT/dpl) is 0.55 pl^-1 at 1.5 L.P.I. and after 4.0 L.P.I. all cells cease elongating in a plane perpendicular to the leaf surface. The formation of cells proceeds exponentially up to 3.0 L.P.I. and about this time cell divisions stop in all parts of the lamina. The mean relative rate of cell formation (dlnC/dpl) at the exponential phase is 1.41 pl^-1, an increase of about 31% per day. At least 27 generations of cells are involved in the process of leaf formation and the generation time was calculated to be 0.5 plastochron or 2.2 days.     

Auxin transport and the regulation of petiole abscission in cotton (Gossypium hirsutum L.): A comparison of indol-3yl-acetic acid and phenylacetic acid

Distal applications of indol-3yl-acetic acid (IAA) to debladed cotyledonary petioles of cotton (Gossypium hirsutum L.) seedlings greatly delayed petiole abscission, but similar applications of phenylacetic acid (PAA) slightly accelerated abscission compared with untreated controls. Both compounds prevented abscission for at least 91 h when applied directly to the abscission zone at the base of the petiole. The contrasting effects of distal IAA and PAA on abscission were correlated with their polar transport behaviour-[1-¹⁴C]IAA underwent typical polar (basipetal) transport through isolated 30 mm petiole segments, but only a weak diffusive movement of [1-¹⁴C]PAA occurred.Removal of the shoot tip substantially delayed abscission of subtending debladed cotyledonary petioles. The promotive effect of the shoot tip on petiole abscission could be replaced in decapitated shoots by applications of either IAA or PAA to the cut surface of the stem. Following the application of [1-¹⁴C]IAA or [1-¹⁴C]PAA to the cut surface of decapitated shoots, only IAA was transported basipetally through the stem. Proximal applications of either compound stimulated the acropetal transport of [¹⁴C]sucrose applied to a subtending intact cotyledonary leaf and caused label to accumulate at the shoot tip. However, PAA was considerably less active than IAA in this response.It is concluded that whilst the inhibition of petiole abscission by distal auxin is mediated by effects of auxin in cells of the abscission zone itself, the promotion of abscission by the shoot tip (or by proximal exogenous auxin) is a remote effect which does not require basipetal auxin transport to the abscission zone. Possible mechanisms to explain this indirect effect of proximal auxin on abscission are discussed.     

Fixation patterns of 14C within developing leaves of eastern cottonwood

Summary Individual leaves of eastern cottonwood (Populus deltoides), representing an ontogenetic series from leaf plastochron index 0.0 to 8.0, were fed ¹⁴CO₂ photosynthetically and then harvested at times ranging from 15 to 1440 min. The lamina of each fed leaf was sectioned from tip to base into 5 parts, and each part was quantitatively assayed for ¹⁴C activity. In young leaves, the percentage of the total ¹⁴C fixed (expressed in dpm/mg of dry leaf tissue) was high in the lamina tip and decreased almost linearly toward the base. With increasing leaf age, the percentage of ¹⁴C fixed decreased in the lamina tip and increased in the base. The relative activity in mature leaves was almost uniform throughout the lamina. No differences were detected in the ¹⁴C distribution patterns within leaves over the time series.On the basis of the data presented and of anatomical observations of developing cottonwood leaves, the hypothesis that the precociously mature lamina tip may provide photosynthates to the still-expanding lamina base was shown to be invalid. It is concluded that bidirectional transport in a developing cottonwood leaf results from simultaneous import to the immature basal region and export from the mature tip.     

Import of 14C-Photosynthate by Developing Leaves of Sugarcane

Sink-to-source transition was studied in developing sugarcane (Saccharum interspecific variety L62–96) leaves. Fully-expanded, mature sugarcane leaves were fed ¹⁴CO₂ for 20 minutes, incorporating about 617 Bq. After five hours the leaves of each plant were cut into 1-cm-length segments that were weighed and then placed in scintillation cocktail for counting. All leaves younger than the leaf fed ¹⁴CO₂ imported labeled photoassimilate. Three to four leaves had both importing and non-importing regions within the blade and a distinct transition region between them. A transition region was observed in leaves which had expanded to between 30 and 90 % of final blade length. Radioactivity per gram fresh weight was calculated as a measure of sink strength. Sink strength was greatest in the youngest leaf and declined with leaf age. The results of this study indicate that 1) import of photosynthate by developing sugarcane leaves occurs over a longer span of developmental ages than in dicotyledonous leaves and 2) the actual tissue region undergoing transition within such a leaf can be resolved as narrow zone between the importing and non-importing regions.     

Translocation pathways in the petioles and stem between source and sink leaves of Populus deltoides Bartr. ex Marsh.

Microautoradiography was used to follow the translocation pathways of ¹⁴C-labeled photosynthate from mature source leaves, through the stem, to immature sink leaves three nodes above. Translocation occurred in specific bundles of the midveins and petioles of both the source and sink leaves and in the interjacent internodes. When each of six major veins in the lamina of an exporting leaf was independently spot-fed ¹⁴CO₂, label was exported through specific bundles in the petiole associated with that vein. When the whole lamina of a mature source leaf was fed ¹⁴CO₂, export occurred through all bundles of the lamina, but acropetal export in the stem was confined to bundles serving certain immature sink leaves. Cross-transfer occurred within the stem via phloem bridges. Leaves approaching maturity translocated photosynthate bidirectionally in adjacent subsidiary bundles of the petiole. That is, petiolar bundles serving the lamina apex were exporting unlabeled photosynthate while those serving the lamina base were simultaneously importing labeled photosynthate. The petioles and midveins of maturing leaves were strong sinks for photosynthate, which was diverted from the export front to differentiating structural tissues. The data support the idea of bidirectional transport in adjacent bundles of the petiole and possibly in adjacent sieve tubes within an individual bundle.Abbreviations C central leaf trace - L left leaf trace - LPI leaf plastochron index - R right leaf trace     

Diurnal oscillatory movements of growing leaves of tobacco

The analysis of diurnal oscillatory movements of tobacco leaves was used in the diagnosis of viral infection of plants. The oscillatory helices circumscribed by a growing leaf of a healthy plant were regular, but some deviations, particularly in the transition points, were recorded. In order to make clear the cause of these irregularities of trajectory, the course of elongation of leaf petiole and blade in relation to localization and shift of zones of elongation during ontogenesis was analysed. The present analysis is similar to that described by the author's earlier experiments with pea roots. Oscillatory curves circumscribed by petiole, projected on a horizontal plane, were compared with curves circumscribed by the blade tip. The analysis of the leaves of different ages enabled us to study this process in dependence on growth rate. It was confirmed that oscillations are a result of elongation; the extent of oscillations is quantitatively dependent on the growth rate. An analysis of the zones of growth showed that in petiole the active meristems are localized near to its base while in the leaf lamina they move gradually during the ontogenesis from the apical to the basal part of the leaf blade. Active meristems of young rapidly growing leaves are localized approximately in the middle of the blade while those of old leaves were found in close proximity to the base of the lamina. The growth rate of petiole can be expressed in hundreds of mm per hour (4.8×10^?2 mm h^?1); half of this value was recorded for its apical part. The growth rate of leaf blade was found approximately ten times higher (3.2×10^?1 mm h^?1). The oscillatory movements of growing leaf consists of two integrate components: of oscillations originating in the base of the petiole and of oscillations of leaf blade the centrum of which is localized in the basal third of the blade. The arrangement of the experiments did not enable us to determine to what extent the phototropic response of leaf blade participates in leaf movements. The movements of leaves of an intact plant are evidently affected by rhythmic stem oscillations. Stem is an integral part of a system which participates in the transfer of information in plants.     

Modification of Pattern of Photosynthate Movement within and between Shoots of Vitis vinifera L

During the prebloom and bloom stages, no movement of labeled photosynthates occurred from a shoot of Vitis vinifera L. fed with ¹⁴CO₂, to an adjacent shoot on the same spur. Movement of labeled assimilates into the unfed shoot was induced when this shoot was sprayed with 2.89 × 10⁻³m gibberellic acid during the prebloom stage. During the bloom stage darkening or defoliation of the unfed shoot resulted in the import of labeled photosynthates from the adjacent fed shoot. Similarly, movement of ¹⁴C into an untreated shoot was induced by removing the terminal 7.5 centimeters and deblossoming the fed shoot. During the berry set stage, translocation of labeled photosynthates from a newly exporting leaf was upwards to the shoot tip, but the direction of movement was reversed by removal of the shoot tip or by darkening or removal of the leaves below the fed leaf. Translocation of photosynthates was predominantly basipetal from a fed leaf near the base of the shoot during the berry set stage, but upward movement was induced by darkening or defoliation of the upper part of the shoot.     

Leaf development and phloem transport in Cucurbita pepo: Carbon economy

Summary Net photosynthesis, dark respiration and growth for leaf 5 of Cucurbita pepo L. plants grown under controlled conditions were measured and the data used for an assessment of the changes in carbon balance during growth of the leaf through expansion to maturity. The blade is first capable of net CO₂ fixation when ca. 8% expanded but the initial rapid growth during this period is sustained almost entirely through imported nutrients. When the growth rate starts to decline rapidly the net photosynthetic capacity of the blade begins to increase. This increase is accompanied by an expansion of the intercellular spaces and by decreasing dark respiration measured at night and in dark periods during the day. The blade becomes completely independent of phloem imported nutrients and begins to export excess photosynthate when the phase of rapid decrease in relative growth rate is almost complete at about 45% expansion. Maximum net photosynthesis of ca. 11 mg CO₂ h^-1 dm^-2 is achieved at 70% expansion. The first detectable synthesis of the transport sugars stachyose and raffinose in the blade coincides with the beginning of intralaminar phloem transport from the tip to the base of the leaf. The synthesis of sucrose, the other major transport sugar, is detectable at all stages of leaf development.     

Auxin Transport as Related to Leaf Abscission during Water Stress in Cotton

Plant water deficits reduced the basipetal transport of auxin in cotyledonary petiole sections taken from cotton (Gossypium hirsutum L.) seedings. A pulse-labeling technique was employed to eliminate complications of uptake or exit of ¹⁴C-indoleacetic acid from the tissue. The transport capacity or the relative amount of radioactivity in a 30-minute pulse which was basipetally translocated was approximately 30% per hour in petioles excised from well watered seedlings (plant water potentials of approximately -4 to -8 bars). No cotyledonary leaf abscission took place in well watered seedlings. Plant water potentials from -8 to -12 bars reduced the transport capacity from 30 to 15% per hour, and although the leaves were wilted, cotyledonary abscission did not increase appreciably at these levels of stress. The threshold water potential sufficient to induce leaf abscission was approximately -13 bars and abscission increased with increasing stress while the auxin transport capacity of the petioles remained relatively constant (15% per hour). The basipetal transport capacity of well watered petioles tested under anaerobic conditions and acropetal transport tested under all conditions were typically less than basipetal transport under the most severe stress conditions. Cotyledonary abscission took place during and 24 hours after relief of stress with little or no abscission taking place 48 hours after relief of stress. Although the water potential returned to -4 bars within hours after rewatering the stressed plants, partial recovery of the basipetal transport capacity of the petioles was not apparent until 48 hours after rewatering, and at least 72 hours was required to return the transport capacity to near normal values. These data support the view that decreased levels of auxin reaching the abscission zone from the leaf blade influence the abscission process and further suggest that the length of time that the auxin supply is maximally reduced is more critical than the degree of reduction.     

Translocation and incorporation of 14C into the petiole from different regions within developing cottonwood leaves

Summary The ability of a developing cottonwood (Populus deltoides Bartr.) leaf to export ¹⁴C-labeled assimilates begins at the lamina tip and progresses basipetally with increasing LPI. This progression indicates that portions of leaves function quasi-independently in their ability to export ¹⁴C-photosynthate. Although most of the exported radioactivity was recovered in the petiole as water-80% alcohol-soluble compounds, there was also substantial incorporation into the chloroform and insoluble fractions. This observation indicates that assimilates translocated from the lamina are used in structural development of the petiole. Freeze substitution and epoxy embedding were used to prepare microautoradiographs for localization of water-soluble compounds. Radioactivity was found in all cell types within specific subsidiary bundles of the petiole. However, radioactive assimilates appeared to move from the translocation pathway in the phloem toward active sinks in the walls of the expanding metaxylem cells. Translocation in the mature xylem vessels was not observed.     

Direct somatic embryogenesis fromBegonia gracilis explants

Direct somatic embryogenesis ofBegonia gracilis was achieved from microcultured laminar segments and petioles on Murashige and Skoog medium with 0.5 mg 1^–1 kinetin and 2% coconut water. Somatic embryos were obtained with greater frequency from petiole explants than from leaf blade sections. Under red light (45 mol m^–2 s^–1), approximately 80% of the petiole explants successfully produced somatic embryos but only 30% of the leaf blade sections responded. However, somatic embryos were significantly more abundant on responding lamina explants (60–70 embryos/leaf section) than on petioles (40–50 embryos/petiole). These trends were similar for explants kept in the dark, but overall production was lower. Somatic embryos were produced more quickly (5 weeks) from petioles than from lamina explants (8 weeks). The somatic embryos germinated to produce plantlets and subsequently shoot cultures with the same appearance as the parental clone.Abbreviations BA benzyladenine - MS Murashige and Skoog (1962) - NAA naphthalene acetic acid - SE somatic embryo     

ORGANIZATION AND ONTOGENY OF THE VASCULAR SYSTEM IN THE PETIOLE OF EASTERN COTTONWOOD

The topologic arrangement of petiolar bundles varies within the length of the cottonwood petiole. Each petiolar bundle is formed by the subdivision and aggregation of acropetally differentiating subsidiary bundles in a predictable pattern. The subsidiary bundles provide vascular continuity between the stem and specific portions of the leaf lamina. Spot-labeling of individual veins with ¹⁴CO₂, freeze substitution, and microautoradiography were used to establish the relation between the secondary veins of the lamina and the vasculature of the petiole. Within the petiole vasculature each subsidiary bundle was continuous with a specific portion of the lamina and seemed to have a separate function. Subsidiary bundles continuous with the central leaf trace were closely related functionally to the tip region of the lamina, while the subsidiary bundles continuous with the lateral leaf traces were functionally related to the middle and basal portions of the lamina.     

AN ANALYSIS OF LEAF ELONGATION IN XANTHIUM PENSYLVANICUM PRESENTED IN RELATIVE ELEMENTAL RATES

Maksymowych , Roman . (Villanova U., Villanova, Pa.) An analysis of leaf elongation in Xanthium pensylvanicum presented in relative elemental rates . Amer. Jour. Bot. 49(1): 7–13. Illus. 1962.—Xanthium plants were grown vegetatively, and leaves, whose developmental stages were specified by a previously described leaf plastochron index (L.P.I.), were marked with India ink along the midrib and photographed during 3 successive days. The relative elemental rates of elongation, d(dX/dpl)/dX were estimated during the whole course of development. The pattern of elongation was not constant but was changing with increasing plastochron age of the leaf. The elements of a young leaf of L.P.I. 0.75 elongated with a constant relative rate. In older leaves, the d(dX/dpl)/dX values were progressively declining toward the tip of the lamina. After L.P.I. 6.3 the only increment in length was due to the elongation of the elements of the petiole. The pattern of growth distribution is discussed in terms of relative elemental rates with respect to cell division and cell elongation in various portions of the lamina and is correlated with the basipetal trend of tissue differentiation in the developing Xanthium leaf.     

Effect of ethylene treatment on transport and metabolism of indole-3-butyric acid in citrus leaf midribs

Transport and metabolism of radiolabeled indole-3-butyric acid (IBA) were studied in midrib sections of Cleopatra mandarin (Citrus reticulata Blanco) and compared to that of indole-3-acetic acid (IAA). Exogenous IBA was metabolized by the midribs to a polar compound, probably an ester conjugate. Ethylene pretreatment of the midribs reduced their capacity to metabolize IBA by ca. 70% as compared to air pretreatment. IBA transport capacity in the leaf midribs was ca. two times greater in the basipetal direction than the acropetal. The basipetal transport capacity of ³H-IBA was lower than that of ¹⁴C-IAA (ca. 24% and 39% of the uptake, respectively). While ethylene treatment reduced basipetal transport of IAA by ca. 70% it did not affect the transport of IBA. Most of the transported label was found as free IBA, but the reduction of IBA conjugation by ethylene treatment did not affect the transport capacity.     

Phloem Unloading in Developing Leaves of Sugar Beet : I. Evidence for Pathway through the Symplast

Physiological and transport data are presented in support of a symplastic pathway of phloem unloading in importing leaves of Beta vulgaris L. (`Klein E multigerm'). The sulfhydryl reagent p-chloromercuribenzene sulfonic acid (PCMBS) at concentration of 10 millimolar inhibited uptake of exogenous [¹⁴C]sucrose by sink leaf tissue over sucrose concentrations of 0.1 to 5.0 millimolar. Inhibited uptake was 24% of controls. The same PCMBS treatment did not affect import of ¹⁴C-label into sink leaves during steady state labeling of a source leaf with ¹⁴CO₂. Lack of inhibition of import implies that sucrose did not pass through the free space during unloading. A passively transported xenobiotic sugar, l-[¹⁴C]glucose, imported by a sink leaf through the phloem, was evenly distributed throughout the leaf as seen by whole-leaf autoradiography. In contrast, l-[¹⁴C]glucose supplied to the apoplast through the cut petiole or into a vein of a sink leaf collected mainly in the vicinity of the major veins with little entering the mesophyll. These patterns are best explained by transport through the symplast from phloem to mesophyll.     

Untersuchungen zur funktionellen Anatomie des Leitgewebesystems im Blatt von Pelargonium zonale

Summary The petiole of Pelargonium zonale is traversed by 17 bundles, whose arrangement and form are typical for this plant. The bundles of the petiole are connected with the conducting system of the axis and with the main nerves by a system of phloem anastomoses in the leaf base and in the junction between the petiole and the leaf blade (Fig. 2). The anatomical findings were confirmed and extended by a study of the translocation of K-fluorescein and ¹⁴C. It could be shown that the metaphloem of the central petiole bundle is composed of phloem subunits, each of which is connected with the phloem of one certain main nerve only (Fig. 4). Accordingly, if fluorescein or ¹⁴CO₂ is applied to one main nerve, the dye or ¹⁴C-material is translocated exclusively in a small phloem area of the central bundle. Autoradiograms of the petioles indicate that the ¹⁴C-labelled assimilates (sucrose, glucose, fructose and amino acids) are translocated exclusively in the phloem. A lateral movement of the labelled material within the petiole was not observed. The metaphloem of the central petiole bundle of Pelargonium zonale revealed a functional organization of phloem subunits.

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Teil einer Dissertation unter der wissenschaftlichen Leitung von Prof. Dr. J. Willenbrink.     

Movement and Endogenous Levels of Abscisic Acid during Water-Stress-induced Abscission in Cotton Seedlings

In an effort to investigate possible involvement of abscisic acid (ABA) in foliar abscission processes, its movement and endogenous levels were examined in cotyledons taken from cotton seedlings (Gossypium hirsutum L.) subjected to varying degrees of water deficit, a condition which initiates leaf abscission. Using a pulse-labeling technique to avoid complications of uptake and exit from the tissue, ABA-1-¹⁴C movement was observed in both basipetal and acropetal directions in cotyledonary petioles taken from well watered, stressed, and rewatered plants. The label distribution patterns obtained after 1 and 3 hours of transport under all situations of water supply were diffusive in nature and did not change when tested under anaerobic conditions. The transport capacity of the petioles ranged from 3.6 to 14.4% ABA-1-¹⁴C transported per hour at estimated velocities of 0 to 2 millimeters per hour. Comparison of basipetal and acropetal movement indicated a lack of polarity under all conditions tested. These low transport capacities and slow velocities of movement, when compared to the active transport systems associated with auxin movement, as well as the lack of anaerobic effects and polarity, suggest that ABA movement in cotton cotyledonary petiole sections is facilitated by passive diffusion. Increases in free and bound ABA in the lamina with increased water stress did not correlate with patterns of cotyledonary abscission. Thus, no evidence was found to suggest that ABA is directly involved in stress-induced abscission processes.     

CELL DIVISION AND CELL ELONGATION IN LEAF DEVELOPMENT OF XANTHIUM PENSYLVANICUM

Maksymowych, Roman. (Villanova U., Villanova, Pa.) Cell division and cell elongation in leaf development of Xanthium pensylvanicum. Amer. Jour. Bot. 50(9) : 891–901. Illus. 1963.—Cell division in different parts of the lamina and cell enlargement of the upper epidermis and palisade mesophyll were studied in vertical and horizontal planes during the entire period of growth. The leaf plastochron index (L.P.I.) was used for designation of developmental stages of the leaf. From cell-length data and measurements of cell area the absolute rates of elongation (dX/dpl) and relative rates of elongation (dlnX/dpl) were calculated. The increase in number of cells in the early plastochrons is exponential and cell division stops at about L.P.I. 3.0. Divisions cease first at the tip and last in the basal lobes of the leaf, indicating a basipetal trend of this process. Cells are elongating while division is in progress, though this elongation proceeds at low rates and for a limited time. Palisade cells elongate in the vertical plane at higher rates and at least 1 plastochron sooner than the upper epidermis. The latter cells, however, expand in area with higher absolute and relative rates, and about 2 plastochrons in advance of the palisade mesophyll. The rates are not constant during the whole period of development but are represented by the bell-shaped curves with maximal peaks around L.P.I. 3.0 for the middle portion of the lamina. The increase in volume of the 2 types of cells stops around L.P.I. 5.0, or shortly after. In addition to unequal durations of cellular enlargement, both tissues expand at differential rates, which for the upper epidermis is high in the horizontal plane but low in the vertical plane, while the opposite is true for the palisade mesophyll. It is suggested that palisades and spongy mesophyll are separated and intercellular spaces formed during the course of development because of the greater rate of expansion in area of the upper epidermis.