Structure and development of the ovule and seed in Aristolochiaceae, with particular reference to Saruma

All members of Aristolochiaceae have anatropous, bitegmic, crassinucellate ovules, which are endostomic except in Saruma and Asarum arifolium where ovules are amphistomic. The outer integument is two cell-layered and the inner integument is three cell-layered. The chalazal megaspore is the functional one. All these conditions appear to be plesiomorphic for the order Piperales, which consists of five families, Aristolochiaceae, Hydnoraceae, Lactoridaceae, Piperaceae and Saururaceae. The embryo sac in Aristolochiaceae is eight-nucleate and corresponds to the Polygonum type; a hypostase is frequently present in this family. The seed coat of Aristolochia s.l., Asarum, Saruma and some Thottea species consists primarily of a two cell-layered testa, and a three cell-layered tegmen. In some species the cells of the outer epidermis become radially elongated, forming reticulate wall thickenings. Cells of the inner layer of the testa have crystals and thickened inner walls. The three layers of the tegmen are tangentially elongated, and become cross fibres at maturity, as fibres of the outer and inner layers are parallel to the seed axis, whereas those of the middle layer are perpendicular to it. This type of seed coat anatomy is synapomorphic for Aristolochiaceae. In addition, the gross morphology of the seed and elaiosome histology are remarkably similar in Asarum and Saruma, thus supporting a sister-group relationship between them. Embryological and seed characters do not supply any synapomorphy that support a close relationship between Aristolochiaceae, Hydnoraceae and Lactoridaceae. Instead, some seed features such as the absence of seed appendages and the collapsed cells of endotesta may indicate a close relationship of Lactoris with Piperaceae plus Saururaceae, although this is the subject of further analysis.     

Asteropeia and Physena (Caryophyllales): A case study in comparative wood anatomy

Previous analyses ofAsteropeia andPhysena have not compared the wood anatomy of these genera to those of Caryophyllales s.l. Molecular evidence shows that the two genera from a clade that is a sister group of the core Caryophyllales. Synapomorphies of theAsteropeia-Physena clade include small circular alternate pits on vessels, presence of vasicentric tracheids plus fiber-tracheids, presence of abaxial-confluent plus diffuse axial parenchyma, and presence of predominantly uniseriate rays. These features are analyzed with respect to habit and ecology of the two genera. Solitary vessels, present in both genera, are related to the presence of vasicentric tracheids. Autapomorphies in the two genera seem related to adaptations byPhysena as a shrub of moderately dry habitats (e.g., narrower vessel elements, abundant vasicentric tracheids, square to erect cells in rays) as compared to alternate character expressions that seem related to the arboreal habit and humid forest ecology ofAsteropeia. The functional significance of vasicentric tracheids and fiber-tracheids in dicotyledons is briefly reviewed in the light of wood anatomy of the two genera.     

Wood anatomy of Gentianaceae, tribe Helieae, in relation to ecology, habit, systematics, and sample diameter

Twenty collections representing one species each ofSymbolanthus andTachia, and 17 species ofMacrocarpaea were studied by means of light microscopy and scanning electron microscopy (SEM). Wood details show that the three genera form a coherent group;Tachia differs from the others in only a few minor characters. Because the species studied form a natural group, wood variations within Helieae offer the basis for correlations and interpretations with respect to habit and ecology. Diameter of stems studied proves to be an important variable that must be taken into account. Correlations with stem diameter include wider vessels in outer wood of wider samples. This would correspond to deeper penetration of reliable water tables by roots of helioid trees or large shrubs. Ray height decreases with increase in stem diameter, an indication of paedomorphosis. Rays of all species are paedomorphic in histology by virtue of relative paucity or even absence of procumbent cells in multiseriate rays. Pseusoscalariform lateral wall pitting of vessels is also a feature characteristic of paedomorphosis. The assemblage of paedomorphic features correlates well with the conclusion, reached by authors who used cladistic methods, that Gentianaceae other than Gentianeae are derived from suffrutescent prennials. The Mesomorphy Ratio, which incorporates three vessel features, correlates with leaf length and with stem diameter. All Helieae are mesophytic, but to various degrees. Septate fiber-tracheids, where present, are typically near vessels and form a substitute for or an addendum to vasicentric axial parenchyma as a mechanism for photosynthate storage. Vestured pits occur on lateral wall pits of vessels of all Helieae, but not on the fibertracheids. Vestured pits show diversity withinMacrocarpaea, a feature of possible systematic significance.     

FLORA OF THE LOWER CRETACEOUS CEDAR MOUNTAIN FORMATION OF UTAH AND COLORADO. PART III: ICACINOXYLON PITTIENSE N. SP

Icacinoxylon pittiense, a new species of angiospermous wood from the Lower Cretaceous Cedar Mountain Formation of Utah is described and compared with similar fossil and modem woods. It is distinguished from other species of Icacinoxylon by its thick-walled fiber-tracheids with their walls making up at least 50% of the total diameter of the cells, conspicuous bordered pits with obliquely crossing extended apertures on both the tangential and radial walls of its fiber-tracheids, scalariform perforation plates with as few as four or greater than 30 bars, transitional opposite to scalariform pitting on its vessel walls, thick-walled ray cells, and distinct sheath or border cells in its rays. Icacinoxylon pittiense is the first species of this genus to be reported from Cretaceous sediments. This wood is of special interest because very few angiosperm woods have been reported from lower Cretaceous strata.     

Systematic and phylogenetic value of wood anatomy in Heteromorpheae (Apiaceae,Apioideae)

The wood anatomy of all four woody genera of the tribe Heteromorpheae (Apiaceae, subfamily Apioideae) has been described and compared, based on 40 wood samples (representing nine species of Anginon, one species of Glia, three species of Heteromorpha and two species of Polemannia). The four genera were found to be relatively similar in their wood anatomy. Helical thickenings on the vessel walls occur in all species investigated and appear to represent an ancestral character state and a symplesiomorphy for the tribes Bupleurieae and Heteromorpheae. Each of four genera has a diagnostically different combination of character states relating to the diameter of vessels, size of intervessel pits, length of fibres, presence and arrangement of banded axial parenchyma, size of rays and ray cells, and presence of septate fibres and crystals in the ray cells. The occurrence of marginal axial parenchyma in Anginon and Glia may be an additional synapomorphy for these taxa. Variation in the wood anatomy of 31 samples from nine species of Anginon is not correlated with habitat (Fynbos or Succulent Karoo Biomes), but instead appears to reflect adaptations to seasonal aridity found in both ecosystems. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 158 , 569–583.     

EMBRYOLOGY AND KARYOMORPHOLOGY OF LACTORIDACEAE

The embryology and karyomorphology of Lactoris fernandeziana, representing the monotypic family Lactoridaceae, were studied in an attempt to clarify its relationships. Embryologically, Lactoris is characterized by a combination of many generalized, plesiomorphic features, which are mostly shared with Magnoliales and partly shared with Laurales and Piperales, and some specialized, apomorphic features including a tenuinucellate ovule, a small nucellus with early disintegrating nucellar tissue, a nonmultiplicative outer integument, an endothelium, and haustorial endosperm. Karyomorphologically Lactoris is confirmed to have 2n = 40 at metaphase, probably as a tetraploid of x = 10, and more or less distinctive features at inter- and prophase. Comparisons based on its embryological and karyomorphological features suggest that Lactoris is not closely related to any other family. Based on evidence from various sources, we hypothesize that an evolutionary line was derived from a common ancestor with Magnoliales, and then diverged into Lactoris, which retains many primitive magnolialean features, and Piperales (and possibly other groups) with more specialized characteristics. Lactoris seems best placed in its own order, Lactoridales, near Piperales.     

Bordered pits in ray cells and axial parenchyma: the histology of conduction, storage, and strength in living wood cells

Bordered pits occur in walls of living ray cells of numerous species of woody dicotyledons. The occurrence of this feature has been minimally reported because the pits are relatively small and not easily observed in face view. Bordered pits are illustrated in sectional view with light microscopy and with scanning electron microscopy in face view for dicotyledonous and gnetalean woods. Bordered pits are more numerous and often have prominent borders on tangential walls of procumbent ray cells, but also occur on radial walls; they are approximately equally abundant on tangential and horizontal walls of upright cells, suggesting parallels to cell shape in flow pathway design. Axial parenchyma typically has secondary walls thinner than those of ray cells, but bordered pits or large simple pit areas occur on some cross walls of parenchyma strands. There is no apparent correlation between the phylogenetic position of species and the presence of borders in ray cells or axial parenchyma. Bordered pits represent a compromise between maximal mechanical strength and maximal conductive capability. High rates of flow of sugar solutions may occur if starch in ray cells or axial parenchyma is mobilized for sudden osmotic enhancement of the conductive stream or for rapid development of foliage, flowers, or fruits. Measurement of the secondary wall thickness of ray cells may offer simple inferential information about the role that rays play in the mechanical strength of woods. © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 153 , 157–168.     

Wood anatomy of Elaeagnaceae, with comments on vestured pits, helical thickenings, and systematic relationships

The secondary xylem of Elaeagnus, Hippophae, and Shepherdia is described and illustrated in detail. Shrubs and small trees of Elaeagnaceae have ring-porous or semi-ring-porous wood with simple perforation plates, vascular tracheids, fiber-tracheids, diffuse or rarely paratracheal axial parenchyma, and uni- or biseriate rays in Hippophae and Shepherdia, but wider rays in Elaeagnus. Walls of vessel elements, especially narrow ones, tracheids, or fiber-tracheids sometimes show helical thickenings; in a few instances these intergrade with small bud-like protrusions associated with pits. Scanning electron microscopy illustrates that small to vestigial vestures are present in all species studied, although nonvestured pits are also common. The analogous nature of vestures and helical thickenings is considered. Comparative wood anatomy suggests a rather isolated position of the family Elaeagnaceae; affinities with Rhamnaceae, Proteaceae, and Thymelaeaceae are discussed.     

Living Cells in Wood 3. Overview; Functional Anatomy of the Parenchyma Network

The very different evolutionary pathways of conifers and angiosperms are very informative precisely because their wood anatomy is so different. New information from anatomy, comparative wood physiology, and comparative ultrastructure can be combined to provide evidence for the role of axial and ray parenchyma in the two groups. Gnetales, which are essentially conifers with vessels, have evolved parallel to angiosperms and show us the value of multiseriate rays and axial parenchyma in a vessel-bearing wood. Gnetales also force us to re-examine optimum anatomical solutions to conduction in vesselless gymnosperms. Axial parenchyma in vessel-bearing woods has diversified to take prominent roles in storage of water and carbohydrates as well as maintenance of conduction in vessels. Axial parenchyma, along with other modifications, has superseded scalariform perforation plates as a safety mechanism and permitted angiosperms to succeed in more seasonal habitats. This diversification has required connection to rays, which have concomitantly become larger and more diverse, acting as pathways for photosynthate passage and storage. Modes of growth such as rapid flushing, vernal leafing-out, drought deciduousness and support of large leaf surfaces become possible, advantaging angiosperms over conifers in various ways. Prominent tracheid-ray pitting (conifers) and axial parenchyma/ray pitting to vessels (angiosperms) are evidence of release of photosynthates into conductive cells; in angiosperms, this system has permitted vessels to survive hydrologic stresses and function in more seasonal habitats. Flow in ray and axial parenchyma cells, suggested by greater length/width ratios of component cells, is confirmed by pitting on end walls of elongate cells: pits are greater in area, more densely placed, and are often bordered. Bordered pit areas and densities on living cells, like those on tracheids and vessels, represent maximal contact areas between cells while minimizing loss of wall strength. Storage cells in rays can be distinguished from flow cells by size and shape, by fewer and smaller pits and by contents. By lacking secondary walls, the entire surfaces of phloem ray and axial phloem parenchyma become conducting areas across which sugars can be translocated. The intercontinuous network of axial parenchyma and ray parenchyma in woods is confirmed; there are no “isolated” living cells in wood when three-dimensional studies are made. Water storage in living cells is reported anatomically and also in the form of percentile quantitative data which reveal degrees and kinds of succulence in angiosperm woods, and norms for “typically woody” species. The diversity in angiosperm axial and ray parenchyma is presented as a series of probable optimal solutions to diverse types of ecology, growth form, and physiology. The numerous homoplasies in these anatomical modes are seen as the informative results of natural experiments and should be considered as evidence along with experimental evidence. Elliptical shape of rays seems governed by mechanical considerations; unusually long (vertically) rays represent a tradeoff in favor of flexibility versus strength. Protracted juvenilism (paedomorphosis) features redirection of flow from horizontal to vertical by means of rays composed predominantly or wholly of upright cells, and the reasons for this anatomical strategy are sought. Protracted juvenilism, still little appreciated, occurs in a sizeable proportion of the world’s plants and is a major source of angiosperm diversification.     

Response of xylem parenchyma by suberization in some hardwoods after mechanical injury

Summary Wound responses of xylem parenchyma by suberization were investigated in some hardwoods by light and electron microscopy. Suberized ray and axial parenchyma cells form a distinct boundary around the wound in all investigated species. Vessels and fibres within and close behind the suberized area appeared more or less occluded; vessels in Fagus, Quercus, and Populus contained suberized tyloses, those in Betula and Tilia contained amorphous and fibrillar deposits. A common mechanism for suberin deposition in the parenchyma cells became evident. Cisternae of the endoplasmic reticulum were apparently involved in suberization. Suberin compounds are extruded by cytoplasmic vesicles, which fused with the plasma membrane, in order to release their content. The suberin layer exhibited the typical lamellated structure; cytoplasmic continuity between suberized cells by plasmodesmata was maintained through the suberin layer. Fagus revealed the most intense suberized area as compared with the other species. Within the reaction zone of Fagus and Quercus, some individual ray and axial parenchyma cells exhibited a subdivision into 2 or 3 compartments prior to suberization. Subdivision was achieved by the formation of a primary wall-like layer. Subsequently, the compartments became individually suberized. Wounding during winter did not induce suberization. Also, samples wounded and kept under water during the vegetation period showed no response. The role of suberization in the effectivity of wound-associated compartmentalization is discussed.     

KLEINODENDRON AND XYLEM ANATOMY OF CLUYTIEAE (EUPHORBIACEAE)

Kleinodendron, a new genus of Euphorbiaceae, was assigned by Smith and Downs to the tribe Cluytieae. A xylem anatomical survey indicates that there are no objections to this placement. Woods of Cluytieae are diverse but may be characterized generally by having pores which average less than 80 μ in diameter and which are well divided between solitary and radial multiple distributions in the same species; simple vessel perforations; alternate intervascular pitting; fiber-tracheids and libriform wood fibers; exclusively uniseriate, or uniseriate and biseriate heterocellular vascular rays in the same species; uniseriate “bridges” linking superposed biseriate ray segments; diffuse, diffuse-in-aggregates, and scanty vasicentric axial parenchyma, sometimes in the same species; and crystal rhomboids. That Microdesmis and Pogonophora diverge sharply from these generalizations in having scalariform vessel perforations and broad vascular rays, is an indication that they may not be closely related to other genera in Cluytieae.     

ANATOMY AND AFFINITIES OF PENTHORUM

The genus Penthorum L. consists of two species of perennial herbs, P. sedoides of eastern North America and P. chinense of eastern Asia. Penthorum has long been considered intermediate between Crassulaceae and Saxifragaceae. An anatomical study of both species was undertaken to contribute to a better understanding of the relationships of these plants. Prominent anatomical features of Penthorum include: an aerenchymatous cortex and closely-spaced collateral vascular bundles of stems; one-trace unilacunar nodes; brochidodromous venation, rosoid teeth bearing hydathodes, and anomocytic stomata of leaves; angular vessel elements with many-barred scalariform perforation plates and alternate to scattered intervascular pits; thin-walled non-septate fiber-tracheids; abundant homocellular erect uniseriate and biseriate rays; and absence of axial xylem parenchyma. In general, Penthorum possesses neither the morphological nor the anatomical synapomorphies which define Crassulaceae, and features shared with Saxifragaceae are largely symplesiomorphous. Thus Penthorum is probably best classified in the monogeneric Penthoraceae.     

Wood structure of Aegiceras corniculatum and its ecological adaptations to salinities

We describe the wood structure of Aegiceras corniculatum and its differences under various soil salinities. This species had diffuse-porous wood with poorly defined growth rings. Vessels which had single perforations occurred abundantly and in multiples and were storeyed. Intervascular pits between contiguous vessels were alternate bordered ones while half-bordered pit-pairs existed between both vessel-ray and vessel-parenchyma. Homogenous xylem rays were multiseriate and uniseriate. Fiber-tracheids with bordered pits often had thinner walls. Xylem parenchyma cells were scant and distributed diffusely and paratracheally. Differences in the structural and quantitative characters of vessels, xylem rays and fiber-tracheids under diverse soil salinities are described. This revised version was published online in August 2006 with corrections to the Cover Date.     

Wood and bark anatomy of Muntingiaceae: A phylogenetic comparison within Malvales s. l.

Quantitative and qualitative data on wood and bark anatomy are given for Muntingia calabura L. and Dicraspidia donnell-smithii Standley. These data are compared with phylogenetic schemes, based on DNA analysis, in which Muntingiaceae belong to the “dipterocarp clade” within Malvales. The data are consistent with this hypothesis, although Muntingiaceae lack pit vestures in vessels, which are seen in the other malvalean families (Cistaceae, Dipterocarpaceae, Neuradaceae, Sarcolaenaceae, Thymeleaceae), and this may represent a loss of pit vestures. All families of the dipterocarp clade agree with both genera of Muntingiaceae in having tracheids as the imperforate tracheary element type (at least ancestrally), although fiber-tracheids also occur in some Dipterocarpaceae and Thymeleaceae. The large size of some malvalean families (with attendant greater diversity in character states) and a paucity of wood studies in those families make for difficulty in comparison of features such as axial parenchyma and ray types with those of Muntingiaceae; character states of these features are consistent with placement of Muntingiaceae in the dipterocarp clade of Malvales. Banded phloem fibers in bark of Muntingiaceae are much like those of other Malvales. Wood of Muntingiaceae is highly mesomorphic according to quantitative vessel features.     

Wood and bark anatomy of Gnetutn gnemon L.

Quantitative and qualitative data are presented for seven collections representing two varieties (unlike in habit) of Gnetum gnemon. Tracheids are present, but abundant and intermixed with them are septate fibre-tracheids rich in starch. Axial parenchyma has been reported only once previously for the species. Axial parenchyma is in strands of 4–10 cells, is rich in starch, is primarily vasicentric (paratracheal) in distribution, less commonly diffuse. About equally common are simple and compound perforation plates; the latter are composed of from two to about ten bordered foraminate perforations, the shape of which may be altered by crowding or coalescence, but is clearly still foraminate. Lateral walls of vessels bear pits that are vestured around pit cavities, not facing the pit membrane. Rays are composed mostly of procumbent cells; the tangential walls bear bordered pits. Crystals, present in ray cells and (rarely) axial parenchyma vary widely in size. Crystalliferous sclereids with layered walls, starch-rich parenchyma, and gelatinous secondary phloem fibres are the main components of bark. Early stages in origin of successive vascular cambia in bark are newly described. When representative conditions are derived from study of large numbers of slides, the classical view that Gnetum vessels are unlike those of angiosperms is supported. Features of Gnetum gnemon wood are discussed in the light of ecology and conductive physiology.     

Myosin,microtubules, and microfilaments: co-operation between cytoskeletal components during cambial cell division and secondary vascular differentiation in trees

The immunolocalisation of unconventional myosin VIII ('myosin') in the cells of the secondary vascular tissues of angiosperm (Populus tremula L. x P. tremuloides Michx. and Aesculus hippocastanum L.) and gymnosperm (Pinus pinea L.) trees is described for the first time and related to other cytoskeletal elements, as well as to callose. Both myosin and callose are located at the cell plate in dividing cambial cells, whereas actin microfilaments are found alongside the cell plate; actin and tubulin are both associated with the phragmoplast. Myosin and callose also localise to the plasmodesmata-rich pit fields in the walls of living cells, which are particularly abundant within the common walls between ray cells and between ray cells and axial parenchyma cells in the phloem and xylem. In those xylem ray cells that contact developing vessel elements and tracheids, myosin, tubulin, actin and callose are localised at the periphery of developing contact and cross-field pits; the respective antibodies also highlight the bordered pits between vessels and between tracheids. The aperture of the bordered pits, whose diameter diminishes as the over-arching border of these pits develops, also houses myosin, actin and tubulin. Myosin, actin and callose are also found together around the sieve pores of sieve elements and sieve cells. We suggest that an acto-myosin contractile system (a 'plant muscle') is present at the cell plate, the sieve pores, the plasmodesmata within the walls of long-lived parenchyma cells, and at the apertures of bordered pits during their development.     

Comparative wood anatomy in Abietoideae (Pinaceae)

This study, which includes 51 species and six genera of subfamily Abietoideae (Pinaceae), assesses the systematic significance of the wood structure in this group. In particular, the presence of normal and traumatic resin canals, the ray structure and the axial parenchyma constitute phylogenetically informative features. Comparative wood anatomy of Abietoideae clearly supports the monophyly of the genera Abies–Cedrus–Keteleeria–Nothotsuga–Pseudolarix–Tsuga, all of which have axial parenchyma with nodular transverse end walls in the regions of growth ring boundaries, crystals in the ray parenchyma and pitted horizontal and nodular end walls of ray parenchyma cells. Axial resin canals support a subdivision of the subfamily into two groups: Abies, Cedrus, Pseudolarix and Tsuga, without axial resin canals, and Keteleeria and Nothotsuga, with axial resin canals and a specific arrangement of traumatic axial resin canals. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 160 , 184–196.     

Differences in the timing of cell death,differentiation and function among three different types of ray parenchyma cells in the hardwood <Emphasis Type="Italic">Populus sieboldii</Emphasis> × <Emphasis Type="Italic">P</Emphasis>. <Emphasis Type="Italic">grandidentata</Emphasis>

Differences in the timing of cell death, differentiation and function among three different types of ray parenchyma cells in the hardwood Populus sieboldii × P. grandidentata which form uniseriate and homocellular rays were examined and clarified. Ray parenchyma cells died within 5 years, and the disappearance of nuclei from ray parenchyma cells did not occur successively from the pith side, even within individual radial cell lines of a given ray. Cell death occurred earliest in contact cells, which were connected to adjacent vessel elements through pits, in the fourth annual ring from the cambium. Cell death occurred next in intermediate cells, which were located within the same cell lines as contact cells but were not adjacent to vessel elements, in the fourth annual ring from the cambium. Finally, isolation cells, which were located within the other cell lines of a given ray, died in the fifth annual ring from the cambium. Secondary wall thickenings in contact cells and intermediate cells were initiated before those in isolation cells in the current year’s xylem. Most starch grains were localized in intermediate cells, and there were more lipid droplets in contact cells and intermediate cells than in isolation cells. In addition, the largest quantities of protein were found in contact cells. Our results indicate that the position within a ray and neighboring short-lived vessel elements might affect the timing of cell death and differentiation and, thus, the function of long-lived ray parenchyma cells in Populus sieboldii × P. grandidentata.     

Comparative palynology and wood anatomy of <Emphasis Type="Italic">Taxodium distichum</Emphasis> (L.) Rich. and <Emphasis Type="Italic">Taxodium mucronatum</Emphasis> Ten.

A comparative study of Taxodium distichum (L.) Rich. and Taxodium mucronatum Ten. was carried out on the basis of pollen morphology and wood anatomy by light and scanning electron microscopy. We describe a detailed analysis of the anatomical characteristics of the wood, including the tracheids, ray parenchyma, axial parenchyma and number of cross-field pits. Palynological characters were also studied to reveal the shape, size and ultrastructure of the pollen grains. These studies give taxonomic support for the recognition of T. distichum and T. mucronatum as two different species.