VASCULAR ANATOMY OF THE NODAL REGION IN POPULUS DELTOIDES BARTR

The ontogeny of vascular bundles in the nodal region of Populus deltoides Bartr. was examined to understand more thoroughly the structure-function relation between leaf and stem. Three vascular traces from the stem independently enter each leaf in the nodal region. At the base of each developing leaf a region was observed in which both bundle size and vascular development was reduced; this region was referred to as the constricted zone. The constricted zone was described quantitatively at 13 locations within the nodal region of a leaf at LPI 5 by determining the number of metaxylem vessels and the total metaxylem vessel area in each of the three leaf traces. A plot of these data showed a distinct minimum value for total metaxylem vessel area within the constricted zone of each trace; the location of this minimum value was referred to as the constriction plane. Each vascular bundle within the nodal region is composed of independent subsidiary bundles that originate within the constricted zone. These bundles provide a direct connection between the leaf lamina and the stem. The node was defined anatomically on the basis of the ontogenetic development of the subsidiary bundles. The node began at the initial exit point of the central trace from the vascular cylinder and extended distally to the constriction plane. This definition allowed us to quantify the limits of each node. The origin of the initiating layer and metacambium was also examined within the nodal region. These precursors of the cambium develop continuously and acropetally from the stem into the leaf. The developmental implications of the constricted zone and the metacambium within the nodal region are discussed with respect to wood formation.     

ESTABLISHMENT OF THE VASCULAR SYSTEM IN SEEDLINGS OF POPULUS DELTOIDES BARTR.

Seven seedlings ranging from 1 to 25 days old were embedded in Spurr's resin and serially sectioned at 1–2 μm. Sectioning extended from well above the apex downward to the hypocotyl base in the 1–day seedlings and to varying levels in the hypocotyl in the older seedlings. Procambial development was analyzed in its entirety for each seedling, and a composite two-dimensional diagram representing the procambial system of a 25-day-old seedling was prepared. Each cotyledon was served by a double-trace, one-half of which was derived from each of two embryonic bundles. The central traces serving the four primary leaves were in turn derived from the four cotyledonary bundles comprising the double traces. The procambial system serving the cotyledons and the four primary leaves approximated a decussate phyllotaxy. The central traces serving the secondary leaves were arranged in a helix that conformed at first to a 1/3 and then to a 2/5 phyllotaxy. Transitions to higher phyllotactic orders were systematic and reproducible, and they occurred in an orderly sequence in both the central and lateral leaf traces. The manner in which leaf traces diverged from parent traces to serve new leaf primordia provided for vascular redundancy. Thus, the entire vascular system was integrated into a highly functional whole.     

ONTOGENY OF MAJOR VEINS IN THE LAMINA OF POPULUS DELTOIDES BARTR

The ontogeny of the major venation in the lamina of Populus deltoides Bartr. leaves was investigated in relation to the development of original procambial bundles, subsidiary bundles, and their derivatives. Serial sections and clearings were used to show that the midrib region is a composite structure consisting of several independent vascular bundles, each of which eventually diverges into the lamina to become a secondary vein. The sequence of events in the ontogeny of major secondary veins is: (1) an original procambial strand develops acropetally and becomes the precursor of the first vascular bundle of the midrib region of the lamina, (2) ground tissue at the forefront of acropetally developing subsidiary procambial bundles differentiates in a wavelike continuum; meristematic regions precede the acropetally developing procambial bundles, (3) discrete subsidiary bundles differentiate in the meristematic regions as they advance acropetally, (4) subsidiary bundles diverge obliquely in the lamina margin giving rise to the secondary veins in a basipetal fashion, and (5) subsequent differentiation and maturation of the secondary veins occurs within the lamina. The original procambial bundles and first-formed subsidiary bundles become the secondary veins of the uppermost portions of the lamina, the next-formed subsidiary bundles become the secondary veins of the middle portions of the lamina, and the last-formed subsidiary bundles become the secondary veins of the lowermost portion of the lamina.     

DEVELOPMENT AND ORGANIZATION OF THE PRIMARY VASCULAR SYSTEM IN POPULUS DELTOIDES ACCORDING TO PHYLLOTAXY

An actively growing cottonwood bud was embedded in epon-araldite and serially sectioned at 2 μm. The sections were analyzed microscopically with the optical shuttle system of Zimmermann and Tomlinson, and all data were quantitatively recorded relative to the apex and to leaf plastochron index (LPI). Analysis of the sections revealed an acropetally developing procambial system organized according to a precise phyllotaxy. Six procambial strands could be recognized and followed long before the leaf primordia that they would enter were evident at the apex. Origin of these strands coincided with developmental events both in the parent trace and its primordium and in the antecedent leaf on the same orthostichy. Once a primordium and its trace attained a certain stage of development, trace bundles began to develop basipetally from the primordium base. These trace bundles appeared to be the earliest progenitors of wood formation in cottonwood. It was concluded that the concept of residual meristem and its corollary, the hypothesis that acropetally developing procambial strands determine the inception sties of new primordia, apply to the cottonwood apex.     

STRUCTURAL CHANGES IN POPULUS DELTOIDES TERMINAL BUDS AND IN THE VASCULAR TRANSITION ZONE OF THE STEMS DURING DORMANCY INDUCTION

Morphological and anatomical changes in shoots of vigorously growing cottonwood plants (Populus deltoides Bartr.) were studied during dormancy induction in 8-hr short days (SD) and in control plants grown in 18-hr long days (LD). Pronounced structural changes occurred in terminal buds after 4 wk and full dormancy was achieved in 7 wk of SD. Leaf expansion ceased after 5 wk of SD as foliage leaves matured to the terminal bud base at leaf plastochron index 0 (LPI 0). Within the bud, total leaf length (lamina + petiole) decreased and stipule length increased progressively each week; thus, the ratio total leaf length/stipule length decreased rapidly, especially at the position of incipient bud-scale leaves LPI - 1 and LPI - 2. These bud-scale leaves were fully developed by wk 6 and were derived from enlarged stipules and aborted laminae. The full complement of primordia within the bud at the start of SD eventually matured as foliage leaves and the first bud-scale leaf (LPI - 1) was initiated immediately following transfer to SD. Acropetal advance of the primary-secondary vascular transition zone (TZ) was associated with leaf maturation. However, it did not advance throughout the entire vascular cylinder as in LD, but only in those leaf traces serving mature leaves beneath the terminal bud. In both LD and SD treatments the same linear relationship was maintained between LPI of the TZ and LPI of the most recently matured leaf; both parameters simultaneously increased in LD and decreased in SD. Thus, the relationship between leaf maturation and advance of the TZ was maintained irrespective of environment.     

LEAF DEVELOPMENT,PHOTOSYNTHESIS, AND C14 DISTRIBUTION IN POPULUS DELTOIDES SEEDLINGS

Rates of net photosynthesis and dark respiration and distribution of C¹⁴ from selected leaves were determined for young cottonwood (Populus deltoides) trees at different stages of development. Four series of five trees—one series for each of four treated leaf positions—were included in the study. Maximum C¹⁴ export occurred when a leaf had just attained maximum size. Lower stem leaves reached maturity quickly and began exporting photosynthate when demands of the young seedling were high. Leaves at higher stem positions matured more slowly, but senescence was also delayed so their effective export life was prolonged. Translocation from a newly exporting leaf was primarily upward to developing leaves and the apex. As a leaf at any one position aged, the translocation pattern gradually shifted from upward to bi-directional and finally to a predominantly downward direction. Photosynthate translocated downward was incorporated into stem wood and roots. Maximum photosynthetic efficiency coincided with the downward shift of C¹⁴ export. Thereafter, net photosynthesis began to decline, at first slowly and then more rapidly. The patterns of photosynthesis, respiration, and C¹⁴ export associated with leaf age all varied according to leaf position on the stem.     

PROCAMBIUM VS. CAMBIUM AND PROTOXYLEM VS. METAXYLEM IN POPULUS DELTOIDES SEEDLINGS

The concept of a procambium-cambium continuum was examined in Populus deltoides by following its development in serially sectioned bud and stem tissues. As in other species, the term cambium is used to refer to that part of the continuum associated with the formation of secondary vascular tissues; i.e., with secondary growth. However, that part of the continuum associated with the formation of primary vascular tissues is subdivided to facilitate interpretation of the consecutive stages of primary xylem differentiation. Thus, the procambium as envisioned by other authors is subdivided into procambium, initiating layer, and metacambium, all of which develop acropetally and in complete continuity. The procambium is derived from the residual meristem in the form of acropetally developing strands and traces. The initiating layer is represented by the first, tangentially separated, periclinal divisions that delineate the position of the prospective cambium. The metacambium is a later stage during which additional periclinally dividing cells unite the initiating layer into a tangentially continuous meristem within a trace bundle. After establishment of the initiating layer, the procambial trace is completely phloem dominated. Protoxylem differentiation begins in an originating center at the base of the leaf primordium and it progresses basipetally to form the protoxylem pole. Cells of the initiating layer do not contribute to the formation of either protoxylem or protophloem. However, those cells of the initiating layer directly opposite the protoxylem pole divide precociously and later differentiate to metaxylem, thus forming a radial file of protoxylem-metaxylem elements. Protoxylem elements of lateral traces are longitudinally continuous with the protoxylem of their parent traces, whereas those of a central trace are longitudinally continuous with the metaxylem of its parent trace. Metaxylem is formed later than protoxylem and it is derived from the metacambium. Metaxylem does not form a continuous system with protoxylem of the same trace because of the different temporal and spatial origins of the two kinds of xylem. Rather, metaxylem is longitudinally continuous with secondary xylem of older traces below. An attempt was made to determine the functional significance of the pattern of protoxylem and metaxylem differentiation in relation to primary and secondary plant development.     

AXILLARY BUD DEVELOPMENT IN POPULUS DELTOIDES. II. LATE ONTOGENY AND VASCULARIZATION

A nearly mature axillary bud of Populus deltoides was embedded in epoxy and serially sectioned at 6 μm. Sectioning extended from the cataphyll tips to a level in the subtending internode about 6 mm below the bud base. Vascular development was followed through the serial microsections and the vascular system was mapped in its entirety from initiation of the original bud traces to termination of the last recognizable leaf trace beneath the bud apex. Each vascular trace was identified as to its origin, its termination within a foliar organ, and its relation to other traces comprising the bud vascular cylinder. Analysis of these data confirmed the procambial patterns found in Part I of this study. Two original bud traces that diverged from the central trace of the axillant leaf gave rise to two pairs of scale traces in quick succession, and these scale traces become the progenitors of all subsequent vascular traces that were perpetuated within the bud. Just before the bud vascular system separated from that of the stem, a third pair of scale traces diverged from the original bud traces; the latter then receded toward the stem to eventually merge with its vasculature. The third pair of scale traces produced a horizontal vascular connection between stem and bud before terminating in the adaxial cataphyll. The vascular system at first conformed with a ½ vascular phyllotaxy when the original bud traces were initiated, progressed through a ⅓ vascular phyllotaxy in the scale trace system, and terminated at the time of sampling with a ⅖ vascular phyllotaxy in the foliage leaf primordia.     

AXILLARY BUD DEVELOPMENT IN POPULUS DELTOIDES. I. ORIGIN AND EARLY ONTOGENY

Origin and early development of axillary buds on the apical shoot of a young Populus deltoides plant were investigated. The ontogenetic sequence of axillary buds extended from LPI –1 (Leaf Plastochron Index) near the apical bud base to LPI –11, the fifth primordium below the bud apex. Two original bud traces diverged from the central (C) trace of the axillant leaf and developed acropetally. During their acropetal traverse the original bud traces gave rise to three pairs of scale traces. All subsequent scale traces, and later the foliar traces, were derived by divergencies from the first two pairs of scale traces. Just before the bud vascular system separated from that of the main axis, a third pair of traces diverged from the original bud traces to vascularize the adaxial scale. Concomitantly, the original bud traces were inflected toward the main vascular cylinder where they developed acropetally and eventually merged with the left lateral trace of the leaf primordium situated three nodes above the axillant leaf; they did not participate in further vascularization of the bud. During early ontogeny a shell zone formed concurrent with initiation of the original bud traces and lay interjacent to them. The shell zone defined the position of the cleavage plane that formed between the axillary bud and the main axis. The axillary bud apex first appeared in the region bounded laterally by the original bud traces and adaxially by the shell zone. Following divergence of the main prophyll traces from the original bud traces, the apex assumed a new position intermediate to the prophyll traces. Ontogenetic development suggested that the axillary bud apex may have been initiated by the acropetally developing original bud traces under the influence of stimuli arising in more mature vegetative organs below.     

THE VASCULAR TRANSITION REGION OF HELIANTHUS ANNUUS. I. BILATERAL AND UNILATERAL PATTERNS OF DIFFERENTIATION

The vascular transition region of Helianthus annuus L. may exhibit either a unilateral or a bilateral differentiation of primary phloem groups. The unilateral type of transition has been previously described but the bilateral type with its two variations is described here for the first time. The three patterns of transition are compared, and the significance of the variability of transition regions is discussed in terms of basic transition types and phylogenetic importance.     

THE VASCULAR TRANSITION REGION OF HELIANTHUS ANNUUS (COMPOSITAE). II. EFFECT OF IONIZING RADIATION

The vascular pattern in the transition region of Helianthus annuus L. may be altered by ionizing radiation in dosages ranging from 650–19,200 R. The alteration of the vascular cylinder is expressed by (1) the failure of normal tracheary elements to mature and (2) by the induction of accessory xylem and/or phloem strands. The anatomical alterations in the vasculature of the transition region are presented and their significance discussed in terms of seedling survival and the mechanisms controlling vascular differentiation.     

THE RELATIONSHIP OF GELATINOUS FIBERS TO WOOD STRUCTURE IN EASTERN COTTONWOOD (POPULUS DELTOIDES)

Wood samples from Populus deltoides Marsh. were used to investigate the relationship of gelatinous fibers to the size and number of other wood elements. An increase in the amount of gelatinous fibers was related to a decrease in diameter and an increase in wall thickness of non-gelatinous fibers. The relative sizes of rays, vessels, and fibers were inter-related, but all of these wood elements decreased in size with an increase in the amount of gelatinous fibers. Apparently environmental conditions controlling the differentiation of gelatinous fibers also influence the development of associated wood elements. Thus, the adverse physical properties of reaction wood must be attributed to all of these structural differences in the wood and not merely to the presence of large numbers of gelatinous fibers.     

STUDIES ON THE LEAF OF POPULUS DELTOIDES (SALICACEAE): MORPHOLOGY AND ANATOMY

Mature field- and growth-chamber-grown leaves of Populus deltoides Bartr. ex Marsh. were examined with light and scanning electron microscopes to determine their vasculature and the spatial relationships of the various orders of vascular bundles to the mesophyll. Three leaf traces, one median and two lateral, enter the petiole at the node. Progressing acropetally in the petiole these bundles are rearranged and gradually form as many as 13 tiers of vascular tissue in the petiole at the base of the lamina. (Most leaves contained seven vertically stacked tiers.) During their course through the midrib the tiers “unstack” and portions diverge outward and continue as secondary veins toward the margin on either side of the lamina. As the midvein approaches the leaf tip it is represented by a single vascular bundle which is a continuation of the original median bundle. Tertiary veins arise from the secondary veins or the midvein, and minor veins commonly arise from all orders of veins. All major veins–primaries, secondaries, intersecondaries, and tertiaries–are associated with rib tissue, while minor veins are completely surrounded by a parenchymatous bundle sheath. The bundle sheaths of tertiary, quaternary, and portions of quinternary veins are associated with bundle-sheath extensions. Minor veins are closely associated spatially with both ad- and abaxial palisade parenchyma of the isolateral leaf and also with one or two layers of paraveinal mesophyll that extend horizontally between the veins. The leaves of growth-chamber-grown plants had thinner blades, a higher proportion of air space, and greater interveinal distances than those of field-grown plants.     

DEVELOPMENT AND ORGANIZATION OF THE SECONDARY VESSEL SYSTEM IN POPULUS GRANDIDENTATA

The apical 22 cm of a dormant, first-year sprout of Populus grandidentata was sectioned serially, and the primary and secondary xylem systems were studied microscopically and graphically reconstructed. A total of 15 nodes was present on the mature stem and 14 foliar primordia in the dormant bud. The vascular traces in the lower portion of the mature stem conformed to a 2/5 phyllotaxy while those of the upper portion and within the dormant bud conformed to a 3/8 phyllotaxy. The 2/5 to 3/8 phyllotactic transition occurred in an extremely precise and systematic two-step pattern: (1) The lateral traces shifted to a new point of origin on the parent central trace, and (2) three new central traces were initiated in sequence by divergences from left-traces. Metaxylem, when followed downward, conformed to the arrangement of the procambial trace system only within one orthostichy. Below this point, the metaxylem components of lateral traces physically separated from those of the protoxylem and continued downward on a new course. Metaxylem vessels produced by the trace cambium originated from a postulated vessel-generating center at the stem-petiole junction. Each metaxylem vessel developing basipetally through the primary body was continuous with a secondary vessel developing basipetally in the secondary body. Because secondary development closed the vascular cylinder, vessels originating from developing leaves or primordia situated at higher levels in the shoot were displaced radially outward when they entered the secondary xyelm. The distribution of vessels in the secondary xylem can therefore be accounted for by a knowledge of the production and distribution of metaxylem vessels in the primary body.