COMPARISON OF EARLY LATERAL VEIN FORMATION IN LINUM USITATISSIMUM LEAVES WITH THEORETICAL NETWORK MODELS

The plastochron age of the Linum leaf that first exhibited lateral leaf vein divergences, the divergent leaf, increased through shoot ontogeny, but the size of the divergent leaf remained constant. There were progressive decreases in the plastochron and relative plastochron rate of leaf elongation, but no significant change in relative chronological rate of leaf elongation, through ontogeny. Thus, divergent leaves of similar sizes occupied different relative positions in the array of leaves on stems of different plastochron ages. These observations are partially consistent with theoretical network model predictions on early leaf vein development. The empirical data of this study suggest additional features of leaf development that should be incorporated into future simulation models for leaf vein development.     

ROLE OF PARENCHYMA IN LINUM USITATISSIMUM LEAF TRACE PATTERNS

A new notation for leaf trace patterns was developed which is consistent with contemporary contact parastichy phyllotaxis notation. New computer-aided methods for generating accurate stem tissue maps were developed. Application of these methods resulted in clarification of the role that parenchyma differentiation plays in delimiting the procambial template for Linum usitatissimum L. stem vasculature through ontogeny. Study of the tissue maps for the various leaf trace patterns exhibited by Linum stems through ontogeny generated a set of observations which permits more rigorous definition of the developmental rules for vascular pattern formation. Long-known geometric principles of phyllotaxis were found applicable to leaf trace patterns.     

RELATIONSHIPS BETWEEN SHOOT GROWTH AND CHANGING PHYLLOTAXY OF RANUNCULUS

Growth of Ranunculus shoots through ontogeny is quantified by techniques utilizing scanning electron microscopy and studies on living plant material. The order of the contact parastichy phyllotaxy in the apical system is related to the relative plastochron rates of growth of the shoot. There is a change in the (2, 3) contact parastichy pattern of vegetative phyllotaxy to a transitional (3, 5) contact pattern which is maintained through sepal production. Formation of a 5(1, 1) whorl of petal primordia establishes a (5, 8) contact pattern with the sepal primordia. Subsequent initiation of stamen primordia, in spiral sequence, results in (5, 8, 13) triple contacts between petal and stamen primordia. The stamen primordia and carpel primordia arrangement is characterized by a (8, 13) contact parastichy pattern of phyllotaxy. Through ontogeny the volume of the shoot apex progressively increases but the shape of the apex, described by a second degree polynomial, remains constant. The plastochron and the relative plastochron rates of radial and vertical displacement of primordia progressively decrease during transition but there is no alteration of the chronological rate of apical expansion. The change in contact parastichy phyllotaxy through ontogeny is interpreted as a change in the relative positions of primordia insertion on the apex resulting from an increase in apical volume and an increased rate of primordia initiation.     

CHANGE IN EPILOBIUM PHYLLOTAXY DURING REPRODUCTIVE TRANSITION

The ontogeny of Epilobium hirsutum grown under natural summer photoperiod in a glasshouse was divided into vegetative, early transitional, transitional, and floral stages. Bijugate phyllotaxy, common to both the vegetative and early transitional stages, is transformed into spiral phyllotaxy during the transitional stage by an initial change in the divergence angle of a single primordium inserted at a unique level on the shoot. Leaf primordia subsequently are inserted in a spiral arrangement in the indeterminate floral shoot apex. The early transitional shoot apical meristem is about 1.5 times the volume of the vegetative meristem but expands at about two-thirds the relative plastochron rate of volume increment of the vegetative meristem. There are progressive decreases in the plastochron and relative plastochron rates of radial and vertical shoot growth through ontogeny. Relative chronological rates of shoot growth, however, are not altered during ontogeny. Spiral transformation results from changes in the relative points of insertion of leaf primordia on the shoot meristem. These changes are accompanied by an increased rate of primordia initiation on a more circular shoot meristem. The change in phyllotaxy during ontogeny is similar to that which was artificially induced by chemical modification of auxin concentration gradients in the shoot apex, with the additional feature that there is an initial increase in the volume of the shoot meristem prior to the natural spiral transformation. Size of the shoot apical meristem, however, appears to have little influence on Epilobium phyllotaxy; but the geometric shape of the meristem is well correlated with bijugate to spiral transformations. This suggests that geometric parameters of the shoot meristem should be considered in theoretical models of phyllotaxy.     

PETIOLE DEVELOPMENT AND XYLEM DIFFERENTIATION IN XANTHIUM REPRESENTED BY THE PLASTOCHRON INDEX

Petiole development and formation of xylem vessels have been investigated in Xanthium leaves from early ontogeny to maturity. Kinetics of growth was presented in terms of absolute and relative elemental rates of elongation. The process of vascularization was assessed by the number of differentiated xylem vessels. The leaf plastochron index (LPI) developed by Erickson and Michelini (1957) was used for designating the various stages of development. An exponential increase in petiole length was observed between the LPIs –3 and +4 indicating a constant relative rate of 0.20 or 20% increase per day. After cessation of lamina elongation at LPI 8, petiole elongation continued for an additional 5 day period, to LPI 9.5. Relative elemental rate analysis revealed that the basipetal pattern of elongation was maintained throughout the leaf development. At a specific plastochron age, the only growth was due to the petiole elongation. Leaves which ceased elongating had not completed their internal development, since the process of xylem formation continued for several plastochrons, or about 8 days. The highest rate of xylem formation was ten vessels per day at LPI 5. On the average, about five xylem vessels differentiated per day in the middle portion of a Xanthium petiole. Mature petioles contained an average of 218 xylem vessels. About 12 canals of schizogenous origin preceeded the development of the vascular tissue.     

Growth of the Stem of Agropyron repens (L.) Beauv

The growth rate of the stem of Agropyron repens (L.) Beauv.begins to decline when the sixth foliage leaf has expanded butthe relative growth rate declines throughout the period betweenthe production of one and ten mature leaves. On an absolutetime scale there is a progressive decline in growth rate ofsuccessively formed stem (node-internode) units. On a plastochronscale the relative growth rate of successive stem units declineswithin the apical region but increases behind the apex. Thedecline in the apical region is related to a decrease in therate of cell division and in the later formed stem units thereis no significant increase in cell number from the time of theirformation by the apex until the internode is initiated duringtheir fourth plastochron. These changes are related to concurrentchanges in the size of the shoot apex and in rates of leaf growth.     

Comparison of Leaf Plastochron Index and Allometric Analyses of Tooth Development in Arabidopsis thaliana

Abstract Two methods of analyses were used to investigate tooth development in serrate (se) mutant and wild-type Columbia-1 (Col-1) Arabidopsis thaliana leaves. There were almost twice as many teeth with deeper sinuses and two orders of toothing on the margins of serrate compared with Columbia-1 leaves. The main objective of this study was to test three hypotheses relative to the source of polymorphism in tooth development: (i) Teeth share similar growth rates and initial sizes, but the deeper teeth are initiated earlier in leaf development. (ii) Teeth share similar timing of initiation and growth rates, but the deeper teeth have a larger initial size. (iii) Teeth share similar timing of initiation and initial sizes, but the deeper teeth have a faster growth rate. Leaf plastochron index (LPI) was used as the time variable for leaf development. Results showed teeth in se were initiated at −27 LPI, 15 plastochrons earlier than those of Col-1. Serrate leaf expansion was biphasic, with the early phase expanding at half the relative plastochron rate of the later phase, which equaled the constant relative expansion rate of Col-1 leaves. Allometric analyses of tooth development obscured the interactions between time of tooth and leaf initiation and the early phase of leaf expansion characteristic of serrate leaves and teeth. Timing of developmental events that allometric analysis obscured can be readily detected with the LPI as a developmental index. Received 25 January 2000; accepted 17 March 2000     

THE GROWTH OF THE STEM TIP OF KALANCHOËU CV. ‘BRILLIANT STAR’

Stein , Diana B. and o . L. Stein . (Montana State U., Missoula.) The growth of the stem tip of Kalanchoë cv. ‘Brilliant Star.‘ Amer. Jour. Bot. 47 (2) : 132—140. Illus. I960.–The purposes of this investigation were (1) to define as clearly as possible the events in the shoot apex and its immediate derivatives during the ontogeny of the shoot; and (2) to determine the changes which occur during the transition from a vegetative to a reproductive meristem. Rate of leaf production in Kalanchoë is basically constant. The rate of leaf growth subsequent to the early primordial state is, however, dependent on the age of the plant and on the environment in which the plant is grown. By keeping these factors constant a correlation can be demonstrated between the size of the youngest visible leaf and the microscopic primordia. Throughout its ontogeny the general architecture of the shoot apex remains essentially the same. Two tunica layers cover the corpus in the vegetative shoot apex, and even in the flowering meristem these 2 layers can be detected. The apex is essentially flat and blends into the adjacent leaf primordia early in the plastochron. About 10 days after flower induction has been started the apex changes its form to a dome, primarily by increased cell division. At the same time the rate of elongation of the youngest internodes increases thus placing the flowering stem tip atop an elongated stem. Axillary development is ultimately responsible for the development of a dichasium.     

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.     

LEAF AND APICAL GROWTH CHARACTERISTICS IN TRITICUM

Leaf initiation rate, leaf primordium growth rates, and apical volume growth rates were determined for seedlings of Triticum aestivum cv. Ramona 50 under controlled environmental conditions. Three leaf primordia are present in the caryopsis, and three more leaves are initiated within the first two weeks after germination with a mean plastochron length of 95.5 hr. Volume growth rates of the apical region were determined on six apices which had six primordia each. The mean radial expansion rate was 0.467/plastochron, and the vertical expansion rate was 0.457/plastochron. The volume expansion rate was 1.393/plastochron. The mean volume doubling time was 0.498 plastochrons or 47.1 hr.     

Rates of corolla growth in tobacco determined with the plastochron index

The growth of vegetative and reproductive shoots of Nicotiana tabacum L. cv. Xanthi is analyzed with the plastochron index to estimate the relationship between corolla growth and time. The plastochron of leaves 9 through 20 declines steadily at each successive node. The flower plastochron increases steadily during the growth of an individual cyme, with the most distal flower to open having the longest plastochron. Variation in the flower plastochron is the result of variation in the rate of flower initiation, not the growth rate of individual flowers. The corolla has an extended phase of approximately constant relative growth in length (between 0.2 · d^–1 and 0.3 · d^–1) until a peak of growth (0.5 · d^–1) 2–3 d before anthesis. Corollas also have periodic peaks and troughs of growth that are low in amplitude (0.1 · d^–1), but persist throughout most of corolla development. The pattern of corolla expansion contrasts strongly with earlier reports of the pattern of tobacco leaf growth.Abbreviations PI plastochron index - PR plastochron ratio - RGR relative growth rate in length The authors thank: Drs. T. Sage and E.G. Williams for the considerable time and space they invested; the members of Dr. R. Wyatt's laboratory for allowing us to use their computer facilities; A. Tull and M. Smith for their care taken in the green-house. This research was supported by U.S. Department of Agriculture grant GAM-89-01056 and by National Science Foundation grant DCB-87-15799.     

SPECIAL PAPER: THE PLASTOCHRON INDEX: A REVIEW AFTER TWO DECADES OF USE

The plastochron index provides a morphological time scale which has proved more reliable than chronological age in studies relating morphological and physiological development of a whole plant or plant organ. Since its inception in 1957, the index has been utilized in a variety of investigations from leaf ontogeny in cottonwood trees to rhizoid cluster initiation in algae. The plastochron index has been extensively used in studies involving source and sink relationships, leaf anatomy, cell differentiation, and primary vascularization. It has been used in investigations of hormonal regulation of plant growth and in studies of the effects of various environmental factors on developmental processes in crops. This paper reviews some of the literature from 1957 to present concerning the development and use of the plastochron index.     

PHOTOPERIOD INDUCED CHANGE IN PHYLLOTAXIS IN XANTHIUM

Photoperiodic floral induction in Xanthium, achieved by subjecting the plants to two long nights, is accompanied by a transient change of the phyllotaxis from the (2, 3) contact parastichy pattern of vegetative plants, to a (3, 5) pattern during the transition. To specify the phyllotaxis, two parameters were estimated from transverse sections of apical buds of control and treated plants: the divergence angle, α, and the plastochron ratio, a. The plastochron ratio decreased progressively during transition from the vegetative to the reproductive state of growth, from a = 1.48 initially to a = 1.15 six days after the beginning of induction. The divergence angle was not altered during the transition. This change in phyllotaxis is interpreted as a change in the relative positioning of leaf primordia on the transitional apex. This transient change appears to be identical with the previously described long-term change of the phyllotaxis of Xanthium brought about by treatment of plants with gibberellic acid.     

CHANGES IN EPILOBIUM PHYLLOTAXY INDUCED BY N-1-NAPHTHYLPHTHALAMIC ACID AND α-4-CHLOROPHENOXYISOBUTYRIC ACID

Preliminary studies establishing relationships between leaf plastochron index and Epilobium hirsutum L. shoot growth provide a method for rigorous selection of plants utilized in experiments designed to test the working hypothesis that endogenous auxin gradient interactions are factors of phyllotactic control in this species. Application of N-1-naphthylphthalamic acid (NPA), an auxin transport inhibitor, to one of the youngest bijugate primordia on the shoot meristem results in increased growth of the treated primordium. Fasciation between the treated primordium and one of the next primordia to be initiated alters relative vertical spacing of primordia. Angular shifts between subsequent primordia result in spiral transformation of Epilobium bijugate phyllotaxy. Application of α-4-chlorophenoxyisobutyric acid (CPIB), an auxin antagonist, to one of the youngest bijugate primordia on the shoot meristem results in decreased growth of the treated primordium that alters both radial and vertical spacing of primordia. This is followed by angular shifts between subsequent primordia resulting in spiral transformation of the bijugate phyllotaxy. Changes in the growth parameters of NPA- and CPIB-treated shoots are similar. Relative plastochron rates of radial and vertical shoot growth of induced spiral shoots are about half those of lanolin paste control shoots, as are the plastochrons and relative plastochron rates of leaf elongation. Treated shoot meristems have eccentricities of 0.5 as compared to bijugate control meristem eccentricities of 0.7. No significant difference is apparent between basal transverse areas of treated and control shoot meristems. The relative chronological rates of growth of treated shoots are not significantly different from those rates of control shoots. Spiral transformation results from changes in relative positions of leaf primordia insertion on the shoot meristem, not from changes in growth of treated shoots. These changes are accompanied by an increased rate of leaf initiation on a more circular shoot meristem. Existing theoretical models of phyllotaxy are discussed in relation to these chemically induced changes of Epilobium leaf arrangement.     

A METHOD FOR DETERMINING PLASTOCHRON INDICES DURING HETEROBLASTIC SHOOT GROWTH

A plastochron is defined as the time interval between two successive recurring events during the growth of plant shoots, such as leaf initiation. The plastochron index (PI) formulated by Erickson and Michelini (1957, American Journal of Botany 44: 297–305) provides a method for determining 1) morphological equivalence in a developmentally variable sample of shoots and 2) rates of development in microscopic tissues and organs, by expressing shoot age as a function of plastochron number. The PI assumes that homologous organs at successive nodes grow exponentially, at equal rates, and the plastochron remains constant. These three conditions are not met in many shoots that exhibit heteroblasty in their plastochron and the growth rate of organs at successive nodes. An alternative computational method for the PI is presented that uses two measurements taken at different times from the same organ during its exponential growth phase. The method does not assume that the PI is a linear function of time. Results of an analysis of cyme internode growth in two races of Arenaria uniflora (Caryophyllaceae) demonstrate that the method proposed is in good agreement with Erickson and Michelini's (1957) method when shoot growth is not markedly heteroblastic. The current method is also used to determine the nonlinear relation between PI and time in a race of A. uniflora that has heteroblastic cyme growth. The results generalize the PI for use in studies of heteroblasty, and for shoots where the relative plastochron rate cannot be directly determined.     

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     

Morphological and growth rate differences among outcrossing and self-pollinating races of Arenaria uniflora (Caryophyllaceae)

Populations of Arenaria uniflora exhibit intraspecific variation in floral size and degree of protandry in association with the evolution of self-pollination. Heterochrony, or a simple change in the absolute timing or rate of developmental events, is proposed as the evolutionary mechanism underlying the origin of the small, self-pollinating flowers from their large, outcrossing progenitors. Inflorescence growth in two autogamous populations and their related outcrossing progenitors was studied to provide the temporal data necessary for testing the hypothesis of heterochrony. All four races showed significant variation in the growth and mature length of inflorescence organs. Inflorescences of selfing races were smaller, and had slower relative growth rates and a two-fold increase in the plastochron relative to outcrossing populations. The large-flowered races were both significantly protandrous. A more detailed growth analysis of flower development in two races indicated that the selfing flowers develop at a slower rate and for a longer duration relative to outcrossing flowers. The implications of these temporal changes in floral ontogeny for the heterochronic origin of self-pollinating floral forms are considered.     

Rates of Cell Division in the Shoot Apical Meristem of Pisum

The relative rates of cell division in different regions ofthe pea shoot apical meristem were obtained by measuring theincrease in the numbers of metaphases following applicationof colchicine to the plants. Absolute values for the rates ofcell division could be calculated since the average rate ofcell division for the whole apex was known. Measurements ofthe rates of cell division were obtained at defined intervalsduring the course of a single plastochron. Within each regionof the apex the rate of cell division did not change more thanabout two-fold throughout the plastochron. There was very littleor no increase in the rate of cell division associated withleaf initiation. The formation of a leaf primordium and thesubsequent growth of the apical dome apparently result fromchanges in the direction of growth rather than changes in therates of growth. Three main regions were discernible withinthe apical meristem: a region with a slow rate of cell divisionin the apical dome, a region of a faster rate of cell divisionat the base of the apical dome and at the site of initiationof procambial strands, and a region of an intermediate rateof cell division in the newly initiated leaf primordium andthe adjacent part of the shoot axis.     

Study of CO2 exchange processes,resistances to carbon dioxide and chlorophyll content during leaf ontogenesis in poplar

Net photosynthetic (P _N) and dark respiration (R _D) rate, stomatal (rs′) and internal (ri′) resistances to carbon dioxide were measured by gas exchange methods on leaves of different ages, expressed in leaf plastochron index units (LPI) for a fast growing poplar cultivar Unal 2. Although the optimal leaf age differs slightly for the different gas exchange parameters, leaf ontogeny is reflected in the same way in these different parameters. MaximalP _N and minimalrs′ and ri′ values were found at LPI between 6 and 10. Chlorophyll concentrations were lowest at LPI lower than 10 although an increase in two steps was found, when leaf age increases up to maturity.