COMPARATIVE ANATOMY OF THE PETIOLE AND INFRAGENERIC RELATIONSHIPS IN JATROPHA (EUPHORBIACEAE)

Anatomical features of the petiole in several species of Jatropha L. (Euphorbiaceae) are presented as evidence in support of infrageneric relationships. A trilacunar 3-trace nodal pattern is typical for the genus. The vascular supply to the stipules is derived from the branching of the two peripheral leaf traces. The number of vascular bundles range from 11 through 9, 7, 5 and 3, and occur in a ring, as free traces, a medullated cylinder, or as U-shaped free traces. The reduction from nine to three bundles is correlated with the gross morphological features while 11, which occurs only in the section Peltatae (Pax) Dehgan & Webster, presents an increase. Reduction in the number of petiolar traces follows the evolutionary advancement of various taxa. This reduction in traces corresponds with south-north distribution of the species and consequential adaptation to colder and more arid climates in Central America and Africa. Smaller leaves, fewer primary veins and fewer vascular traces have resulted as a response to reduced need for water. Presence of dorsal (super-numerary) bundles which supply the petiolar glands in subgenus Jatropha (= Adenoropium Pax) is considered significant, since African taxa of the section (subsection Pubescentes Pax) have retained these bundles despite the loss of petiolar glands. The latter glands are prominent in the South American and Indian species. Sectional lines in the genus can, therefore, be drawn generally on the basis of numerical constancy and relative uniformity in the arrangement of petiolar traces. The continuity of vascular bundles from the stem into the petiole and variations of bundle arrangements are depicted in three-dimensional drawings.     

XYLEM ANATOMY AND HYDRAULIC CONDUCTANCE OF PSILOTUM NUDUM

Psilotum nudum (L.) Beauv. (Psilotopsida) has a simple, vascularized sporophyte with a dichotomously branching aerial axis. The number and lumen diameters of tracheids in the actinostele decrease in each subsequent branch, leading to an approximate halving of the measured hydraulic conductance (K_h) from segment to segment. To understand how the anatomy of P. nudum affects K_h, a biophysical model based on the Hagen-Poiseuille relation was developed that incorporated lumen diameter, tracheid taper, pit cavities, and pit membranes. Using a technique previously developed for ferns, pit membrane resistance was determined by measuring water flow before and after dissolving the pit membranes with cellulase. Measured K_h was in good agreement with K_h calculated with the model after excluding thick-walled late metaxylem tracheids that dye studies showed were nonconducting. Model simulations showed that the approximately 40% overlap observed for tracheids of P. nudum was in the range leading to greatest conductance and that K_h decreased to half for 20% overlap. The model also showed that the pit membranes account for an increasing percentage of total resistance to water flow as the lumen diameter increases. Thus, the removal of such primary wall material and the evolutionary origin of vessels would have substantially increased K_h.     

XYLEM ANATOMY AND HYDRAULIC CONDUCTANCE OF COSTA RICAN BLECHNUM FERNS

Hydraulic conductivities of stems, stipes, and elongate leaf stipes were determined for greenhouse-grown Blechnum (B. fraxineum, B. fragile, B. buchtienii, B. sprucei) and Salpichlaena (S. volubilis) plants collected in tropical rain forests of Costa Rica. Organ conductivity was examined in relation to morphology and tracheid characteristics in order to gain an understanding of factors influencing water flow. Hydraulic conductivity of plant organs was determined by measurement of transpiration rates, leaf areas, and water potential gradients. Erect stemmed species develop larger whole plant water potential gradients than elongate stemmed species for a similar transpiration rate. Elongate leaves develop even smaller water potential gradients for a given transpiration rate. Stems have larger hydraulic conductivities but smaller leaf-specific conductivities (LSCs) than stipes. Small conductivities and small LSCs are associated with short, erect stems. Elongate structures have large conductivities and large LSCs. Of the tracheid characteristics examined, the most important characteristics determining the magnitude of organ hydraulic conductivity are diameter, pit aperture area between tracheids, taper length, and cell length. Large conductivities of S. volubilis climbing leaf stipes are associated with very large-diameter tracheids (some > 200 μm), large tracheid number, exceptionally long tracheids (some > 4 cm), large pit aperture area between tracheids, short tracheid taper, and smooth tracheid lumen walls. Hagen-Poiseuille estimates of hydraulic conductivity range from 1.1 to 3.3 times the measured values. Conductivity of stipes is highly correlated with leaf area supplied by stipes. Conductivities of stems and elongate leaf stipes also correlate with leaf area supplied by these structures. Estimated hydraulic conductivities of field-grown Blechnum and Salpichlaena demonstrate that larger conductivities are associated with larger plants. This study contributes toward our knowledge of fern water relations and extends previous growth form/hydraulic architecture characterizations by providing a more comprehensive comparison of closely related species. In addition, this study provides evidence for the relative importance of tracheid characteristics in determining the magnitude of organ hydraulic conductivity.     

XYLEM ANATOMY OF SOPHORA (LEGUMINOSAE) IN RELATION TO HABIT

Cumbie , B. G., and Dan Mertz . (U. Missouri, Columbia.) Xylem anatomy of Sophora (Leguminosae) in relation to habit . Amer. Jour. Bot. 49(1): 33–40. Illus. 1962.—A comparison of 8 tree, shrub, and herbaceous species of Sophora showed considerable variation in structural organization of the secondary xylem in relation to habit. A reduction in duration of cambial activity has been accompanied by a decrease in conspicuousness of storied structure, more frequent arrangement of vessels in multiples, the formation of narrower rays, and a striking reduction in amount of axial xylem parenchyma. On the other hand, relatively minor changes in size of xylem elements have resulted from decreased cambial activity, as shown by comparing the first-formed secondary xylem elements of woody and herbaceous species. In many structural features, shrubs were found to be more or less intermediate between the tree and herbaceous species, at least in the early stages of development. These observations indicate that the often repeated generalization that the organization of herbaceous stems resembles that of the first year's growth in closely related woody species is not always applicable.     

DEVELOPMENTAL ANATOMY AND ULTRASTRUCTURE OF THE ANT-FOOD BODIES (BECCARIIAN BODIES) OF MACARANGA TRILOBA AND M. HYPOLEUCA (EUPHORBIACEAE)

Macaranga is a common secondary growth tree of S.E. Asia. Nine species possess hollow stems which harbor an ant colony, and also produce food bodies which are eaten by the ants. In return, the ants protect the plant from herbivore damage. The multicellular food bodies of M. triloba (Bl.) Muell. Arg. are developed on the underside of down-turned clasping stipules, while in M. hypoleuca (Reichb. f. and Zoll.) Muell. Arg. they are produced on the abaxial surface of young leaves. Food body cells of both species are very rich in lipid, contain large starch grains, and possess an electron-dense hyaloplasm. It is proposed to name the Macaranga ant-food bodies Beccariian bodies in honor of the Italian botanist Odoardo Beccari who explored S.E. Asia in the late 1800s.     

RELATION OF DWARFMISTLETOE (ARCEUTHOBIUM) TO THE XYLEM TISSUE OF CONIFERS. I. ANATOMY OF PARASITE SINKERS AND THEIR CONNECTION WITH HOST XYLEM

Srivastava , L. M., and K. Esau . (U. California, Davis.) Relation of dwarf mistletoe (Arceuthobium) to the xylem tissue of conifers. I. Anatomy of parasite sinkers and their connection with host xylem. Amer. Jour. Bot. 48(2): 159–167. Illus. 1961.—The anatomy of the sinkers of Arceuthobium infecting 7 species of conifers was studied by the use of serial cross, radial, and tangential sections of the host wood. The sinkers were found to be composed of parenchyma cells only, or of parenchyma cells and tracheary elements, including vessel elements. In all species tracheary cells of the sinkers had direct contacts with the host tracheids of axial and radial systems. Typically the sinkers were associated with rays of the host wood. In some species, the centripetal ends of sinkers were wedged in radially among the axial tracheids of the host, but centrifugally such sinkers were usually found associated with rays. In the region of the host cambium the sinker contained parenchyma cells meristematic in appearance and, in 6 out of 7 species, also mature tracheary elements. The oldest of these elements became stretched and ruptured, a circumstance indicating that growth occurred in the part of the sinker embedded in the host cambium. This growth appeared to be coordinated with that of the host cambium, so that the sinker became embedded in the host xylem and phloem. Radial centripetal penetration of sinkers among differentiating axial tracheids of the host possibly occurred to a limited extent.     

STIPE ANATOMY,WATER POTENTIALS,AND XYLEM CONDUCTANCES IN SEVEN SPECIES OF FERNS (FILICOPSIDA)

Anatomy and water relations were studied for the desert fern Notholaena parryi, as well as six other ferns representing three different orders which occupied xeric as well as mesic habitats. Tracheid number and diameter, and total xylem cross sectional area increased during leaf development for N. parryi; the whole plant conductance (volume flow of water through a stipe divided by the rhizome-to-leaf water potential drop) increased but tended to level off as the leaves matured. The reported occurrences of very steep water potential gradients (about 25 MPa m^–1) in stipes of N. parryi were confirmed. The ferns with the highest whole plant conductances (Alsophila australis, Botrychium dissectum, and Adiantum capillus-veneris) had the largest or greatest number of tracheids. Numerous tracheids in Botrychium dissectum offset a low tracheary conductivity, whereas Marsilea vestita had few tracheids resulting in a low whole plant conductance. Whole plant conductances for the ferns were 2 to 3 orders of magnitude less than those generally observed for angiosperms and 6 orders less than for gymnosperms. However, the relative conductivity (whole plant conductance times stipe xylem length divided by xylem area) was only 5- to 10-fold less than for angiosperms and about the same as for the gymnosperms. Stipe water relations in these ferns are discussed in relation to the evolution of xylem anatomy.     

THE MORPHOLOGY AND RELATIONSHIPS OF DALECHAMPIA SCANDENS (EUPHORBIACEAE)

The circumtropical but preponderantly American genus Dalechampia, comprising nearly 100 species of twining vines (or rarely subshrubs), is strikingly isolated within the Euphorbiaceae because of its distinctive bibracteate inflorescences. There has been considerable taxonomic controversy with regard to the relationships of the genus, and it has been suggested that Dalechampia is allied to the tribe Euphorbieae because of a supposed analogy between its inflorescence and the cyathium in the Euphorbieae. Field and laboratory investigations of the common American species D. scandens, together with a comparative survey of related species, have thrown some light on these problems. The Dalechampia inflorescence seems best interpreted as consisting of a terminal staminate pleiochasium (with part of the lateral branches transformed for nectar production), juxtaposed to a 3-flowered pistillate cyme. The lips of the conspicuous bilabiate involucre are formed by the hypertrophied bracts which subtend the staminate and pistillate cymes. The bisexual inflorescences appear to be distinctly proterogynous, rather than proterandrous, as has been previously suggested. The configuration of the inflorescence—a bilaterally symmetrical pseudanthium—suggests adaptation for crosspollination, but the closing movement of the bracts makes self-pollination probable in the absence of visits by pollinators. The similarity of the Dalechampia inflorescence to the cyathium of the Euphorbieae appears to be entirely superficial, and both reproductive and vegetative data suggest that Dalechampia is related to taxa of tribe Plukenetieae.     

EFFECTS OF TREE AGE ON SECONDARY XYLEM AND PHLOEM ANATOMY IN STEMS OF GREAT BASIN BRISTLECONE PINE (PINUS LONGAEVA)

Xylem and phloem tissue samples were collected from various-aged Great Basin bristlecone pine (Pinus longaeva D. K. Bailey) stems in southern Utah and southeastern California to determine whether the vascular cambia of older trees produce fewer xylem rays, shorter-lived xylem and phloem ray cells, fewer phloem sieve cells, and a thinner phloem. Increment cores were examined to determine whether ‘aged’ cambia produced narrower tracheids that might reduce water translocation. Sapwood thickness was measured and sapwood growth layers were counted on these cores. Regression and Classification and Regression Tree (CART) analyses of sample data found no age-related changes in cambial products. Phloem and xylem production appeared normal at all ages, with no evidence of cambial malfunction.     

RELATION OF DWARFMISTLETOE (ARCEUTHOBIUM) TO THE XYLEM TISSUE OF CONIFERS. II. EFFECT OF THE PARASITE ON THE XYLEM ANATOMY OF THE HOST12

Srivastava , L. M., and K. Esau , (U. California, Davis.) Relation of dwarfmistletoe (Arceuthobium) to the xylem tissue of conifers. II. Effect of the parasite on the xylem anatomy of the host. Amer. Jour. Bot. (48(3): 209–215. Illus. 1961.—The changes in the xylem anatomy induced by dwarfmistletoe infection were studied in 7 coniferous species. The most pronounced abnormalities are observed in the shape and size of the infected rays. Because of the presence of parasite tissue, the rays assume a hypertrophied appearance; moreover, they fuse to form large composite rays. The union of rays involves intrusive growth of ray cells and displacement of fusiform initials. Some division of fusiform initials also occurs. Rays may increase in number and they may contain more host cells than normal rays. Axial tracheids in infected host woods differ more or less strongly from those of noninfected woods. They may be shorter, wider, and more irregular in shape than the axial tracheids in healthy wood. The samples of xylem from infected pines had a larger number of resin canals than those from healthy trees. Resin canals were also found in infected Tsuga, which normally lacks these structures.     

THE SYSTEMATICS AND ANATOMY OF GONGYLOCARPUS (ONAGRACEAE)

The unique fruiting structures of the closely related, principally Mexican, monotypic genera Gongylocarpus and Burragea (Onagraceae) compelled a detailed anatomical and cytological investigation of these plants which led to the conclusion that they should be included in a single genus, Gongylocarpus. Gongylocarpus fruticulosus (Burragea), endemic to two adjacent islands off the west coast of Baja California, is divided into two subspecies, subsp. fruticulosus and subsp. glaber. The vegetative and floral anatomy, including wood anatomy, of both species is described. The fruits of these two species grow into the stem by meristematic activity during the course of ontogeny, the ovaries in the mature flower being superficial and sessile in the leaf axil. There is no pedicel associated with the flower, but only a branch gap. Meristematic tissue at the base of the locules divides rapidly at a relatively late ontogenetic stage, the ovaries growing downward into the stem and crushing the pith. The mature, heavily sclerenchymatous fruits are located wholly within the stem, and in G. fruticulosus they are aggregated into long chains. Both species have a gametic chromosome number of n = 11, a characteristic otherwise unknown in the tribe Onagreae but shared with other generalized groups in the family. Taken together with other features, this suggests a primitive position within the tribe for Gongylocarpus.     

TYPES OF CAMBIAL ACTIVITY AND WOOD ANATOMY OF STYLIDIUM (STYLIDIACEAE)

Three types of cambial activity, two hitherto unreported, are described for Stylidium. The four species of sect. Rhynchangium of subgenus Nitrangium have woody cylinders in upright stems. In these a cambium formed beneath the endodermis produces a determinate quantity of fibers, vessel elements, and interxylary phloem strands toward the inside but no derivatives toward the outside; this was correctly reported by Van Tieghem and Morot (1884a) but doubted by subsequent workers. The same species have lignotubers in which a cambium produces contorted xylem (mostly vessels) to the inside, phellem toward the outside. In S. glandulosum and S. laricifolium a cambium formed beneath the endodermis produces an indeterminate quantity of xylem (fibers and vessel elements) and interxylary phloem toward the inside, nothing toward the outside. The xylem is rayless and lacks axial xylem parenchyma. These three modes of cambial activity represent innovations within Stylidiaceae. The family has a wholly herbaceous ancestry if one can judge from the total lack of cambial activity in vascular bundles.     

ONTOGENY AND ANATOMY OF THE FLOWER OF LIMNOCHARIS FLAVA (BUTOMACEAE)

The flowers of Limnocharis flava (L.) Buch. are borne in an indeterminate umbel and each consists of three sepals, three yellow petals, and about 18 carpels surrounded by numerous stamens and staminodia. The androecium is centrifugally developed, and the last-formed members are staminodial; it is supplied by branching vascular systems. Carpels arise almost simultaneously, and a prominent residual floral apex remains. The carpels are partially conduplicately closed and are also primitive in possessing laminar placentation and in lacking differentiation of a style. The gynoecium is essentially apocarpous, but there are slight fusions of adjacent carpels near their ventral margins where they are attached to the receptacle. It is suggested that the Limnocharis flower is the most primitive in the family.     

XYLEM ANATOMY,WATER FLOW,AND HYDRAULIC CONDUCTANCE IN THE FERN CYRTOMIUM FALCATUM

Xylem anatomy and water relations were studied in holly fern (Cyrtomium falcatum, Aspidiaceae) to determine the details of the pathway for water flow through an entire plant and the influence of tracheid number and lumen diameter on water flow. Each leaf has two adaxial traces and an abaxial trace, which are supplied by diarch adventitious roots attached to the dictyostele of the rhizome near the leaf base. Anatomical observations and dye experiments showed that each adaxial bundle vascularizes the approximately seven pinnae on its side of a leaf. An abaxial bundle is intermittently connected to an adaxial bundle as well as other abaxial bundles, forming a minor vascular pathway between the bundles of the leaf axis. Changes in both number and diameter of tracheids result in an acropetal decrease in hydraulic conductance per unit length along the rachis, although tracheid number locally increases when the trace for a pinna is produced in an adaxial bundle. Water flow was determined from the transpiration distal to the point in question or by forcing a solution through an axis with applied pressure. The water potential gradient along the plant axis was quite constant, indicating that hydraulic conductance per unit length varied with leaf area to be supplied. About 40% of the overall water potential drop occurred from the rachis into the pinnae, which reflected factors controlling water potential gradients in the lamina and not a very low conductance in the petiolule xylem. Hydraulic conductances calculated using the Hagen-Poiseuille equation and tracheid diameters were generally double those of measured conductances. Since the values tended to vary by a constant factor, tracheid number and diameter may largely control water flow in the xylem.     

HYDROCARBONS FROM PEDILANTHUS MACROCARPUS (EUPHORBIACEAE) OF BAJA CALIFORNIA AND SONORA,MEXICO

Pedilanthus macrocarpus (Euphorbiaceae) from Baja California and Sonora, Mexico is a desert xerophyte rich in polyisoprenes and other highly reduced hydrocarbons. The latex was analyzed for rubber content and found to contain 6–10% rubber by fresh weight. Four geographically distinct populations were analyzed for alkane content and the normal alkane range was shown to be C₂₇H₃₆ to C₃₅H₇₂. The different populations show significant quantitative and qualitative variation in their alkane content. Phytochemical findings suggest that the hydrocarbon chemistry can be useful in understanding the evolution of Pedilanthus macrocarpus in Baja California.     

ONTOGENY OF THE COTYLEDONARY REGION OF CHAMAESYCE MACULATA (EUPHORBIACEAE)

Development of the cotyledonary region in Chamaesyce maculata is described from germination of the seed through formation of the dense mat of branches which characterize this common weed. The cotyledonary node is trilacunar with split-lateral traces. Epicotyl development is limited to a pair of leaves (“V-leaves”) inserted directly above and decussate to the cotyledons. The two V-leaves are also vascularized by three traces and insertion of these traces relative to the vasculature at the immediately subjacent cotyledonary node is asymmetrical; four of the six V-leaf traces arise on one side of the intercotyledonary plane and two arise on the opposite side. Further shoot development is limited to lateral branches that develop sequentially from cotyledonary axillary buds, and then from de novo buds which arise at bases of previously developed lateral branches. The first axillary bud to develop is located on that half of the seedling which supplies the V-leaves with four traces. Development or insertion of the third and fourth branches (first and second de novo branches) relative to the first and second (cotyledonary) branches occurs in two patterns, termed cis and trans. Subsequent de novo branches generally develop between the two most recently developed branches on that half of the seedling, gradually forming a large branch plexus at the cotyledonary region. In mature robust specimens, however, the sequence of lateral branch development may become irregular. Structure of the cotyledonary region of C. maculata does not readily support widely held concepts of homology between the pleiochasium of Euphorbia subgenus Agaloma and the lateral branch system of Chamaesyce.     

POLLINATION OF DALECHAMPIA MAGNOLIIFOLIA (EUPHORBIACEAE) BY MALE EUGLOSSINE BEES

Flowers of Dalechampia magnoliifolia in eastern Peru were pollinated primarily by male Eulaema meriana and E. cingulata, which collected fragrance from the secretory gland borne in the staminate cymule. The fragrance contains carvone oxide, benzyl acetate, limonene, α-pinene, myrcine, β-pinene/camphene, and carvone. Trigona cf. pallens was the most abundant visitor, but was primarily a pollen thief. The rate of contact with the stigmas by Trigona, and hence the bee's efficacy as a pollinator, was highly variable and was determined by variation in the separation of anthers and stigmas. Dalechampia magnoliifolia co-occurred with a congeneric species, D. cissifolia, but did not share pollinators with it.