Water-storing and Cavitation-preventing Adaptations in Wood of Cacti

Ancestral cacti presumably had abundant, fibrous, heavily lignifiedwood, similar to that present in the relictual, leaf-bearinggenus Pereskia. During the evolutionary radiation of the subfamilyCactoideae, diverse types of bodies and woods arose. Severalevolutionary lines have retained an abundant, fibrous wood:all wood cells, even ray cells, have thick lignified walls,and axial parenchyma is only scanty paratracheal. Aside froma diversity of vessel diameters, there seems to be little protectionagainst cavitation during water-stress, and little water-storagecapacity. This strong wood permits the plants to be tall andto compete for light in their tree-shaded semi-arid habitats.In other evolutionary lines, the wood lacks fibres, and almostall cells have thin, unlignified walls. Vessels occur in anextensive matrix of water-storing parenchyma, and tracheidsare also abundant, constituting over half the axial tissue insome species. There is excellent protection against cavitation,but little mechanical support for the plant body; however, theseplants are short and occur in extremely arid, unshaded sites.Scandent, vinelike plants of two genera produce a dimorphicwood—while their shoots are extending without externalsupport, they produce fibrous, lignified wood, but after leaningagainst a host branch, they produce a parenchymatous, unlignifiedwood.Copyright 1993, 1999 Academic Press Cactaceae, cactus, water-stress, wood, evolution, xylem     

Hydraulic architecture and the evolution of shoot allometry in contrasting climates

We used pairs of congeneric shrub species from contrasting habitats to test for repeated evolutionary divergence in leaf-stem allometry and shoot hydraulic architecture in response to water availability. Allometric relationships and mean ratios between leaf size (individual and total area and mass per shoot) and stem cross-sectional area were compared between habitats using six species pairs representing three genera (Arctostaphylos, Baccharis, Ceanothus). We measured correlations among evolutionary changes in allometric, morphological, and physiological traits using phylogenetic independent contrasts. Allometric analysis revealed habitat differences: slopes were homogeneous among species (=1.46), but the more mesic-adapted species generally supported more leaf area at a common stem cross-sectional area. Reducing bivariate allometry to a ratio obscured this pattern because ratios varied with stem size, which was unrelated to habitat. Mean individual leaf size also was not correlated with either water availability or leaf-stem allometry. Stem hydraulic conductivity was generally lower in the xeric-adapted species of each pair, and its evolution mirrored changes in shoot allometry. This study provides evidence for repeated evolutionary divergence in shoot allometry and hydraulic architecture associated with water availability and demonstrates the importance of shoot allometry to water relations, independent of leaf size.     

WOOD ANATOMY AND RELATIONSHIPS OF LACTORIDACEAE

Mature wood of Lactoris, not previously available for study, reveals ten distinctive characters: vessels with simple perforation plates; vessels in pore multiples; vessel-to-axial parenchyma pits scalariform or transitional, vessel-to-vessel pits alternate; fiber-tracheids with vestigial pits; fiber-tracheids, vessels, and axial parenchyma storied; axial parenchyma vasicentric scanty; axial parenchyma either not subdivided or, if subdivided, with thin nonlignified walls between the cells (like the septa in septate fibers); rays wide and tall, little altered during ontogeny; ray cells upright; and ray cells taller adjacent to fascicular areas. All of these features occur in woods of Piper and other Piperaceae. The systematic position of Lactoris is therefore reassessed. Evidence available to date is consonant with placement of Lactoridaceae in Piperales, in which it would be more primitive than Piperaceae or Saururaceae. Features cited as evidence for alternative placements of Lactoridaceae are reviewed.     

Structure-function relationships in highly modified shoots of cactaceae

BACKGROUND AND AIMS: Cacti are extremely diverse structurally and ecologically, and so modified as to be intimidating to many biologists. Yet all have the same organization as most dicots, none differs fundamentally from Arabidopsis or other model plants. This review explains cactus shoot structure, discusses relationships between structure, ecology, development and evolution, and indicates areas where research on cacti is necessary to test general theories of morphogenesis. SCOPE: Cactus leaves are diverse; all cacti have foliage leaves; many intermediate stages in evolutionary reduction of leaves are still present; floral shoots often have large, complex leaves whereas vegetative shoots have microscopic leaves. Spines are modified bud scales, some secrete sugar as extra-floral nectaries. Many cacti have juvenile/adult phases in which the flowering adult phase (a cephalium) differs greatly from the juvenile; in some, one side of a shoot becomes adult, all other sides continue to grow as the juvenile phase. Flowers are inverted: the exterior of a cactus 'flower' is a hollow vegetative shoot with internodes, nodes, leaves and spines, whereas floral organs occur inside, with petals physically above stamens. Many cacti have cortical bundles vascularizing the cortex, however broad it evolves to be, thus keeping surface tissues alive. Great width results in great weight of weak parenchymatous shoots, correlated with reduced branching. Reduced numbers of shoot apices is compensated by great increases in number of meristematic cells within individual SAMs. Ribs and tubercles allow shoots to swell without tearing during wet seasons. Shoot epidermis and cortex cells live and function for decades then convert to cork cambium. Many modifications permit water storage within cactus wood itself, adjacent to vessels.     

三种阔叶树雌雄异株木材结构的比较

在光学显微镜下较详细地比较了重阳木、大叶刺篱木和红毛丹,雌雄异株木材的结构。结果表明：同一种树木两性间木材结构总体上是很相似的,但有的定量特征差异显著甚至极显著。３个种的树木中,雌株木纤维壁腔比值较雄株的大,而分膈纤维和纤维管胞数量较小。雌株木材结构特点可能表明其木材具较强的机械支持能力,利用于果实的支持 。     

Angiosperm wood evolution and the potential contribution of paleontological data

Wood anatomy is often viewed as a source of independent data that may be used to assess evolutionary relationships among angiosperms. Comparative anatomical studies document suites of correlated characters that have been interpreted as general evolutionary trends, of which several have been asserted to be irreversible. Paleobotanical data summarized by Wheeler and Baas provide broad chronological corroboration of some wood anatomical trends, such as evolution from scalariform to simple perforation plates and long to short vessel elements. However, the focus on general evolutionary trends rather than on analyzing character distribution patterns in a cladistic phylogenetic context obscures a more detailed understanding of the evolution of wood anatomical features. Patterns of character evolution, including the assertions of irreversibility, need to be tested through cladistic analyses. In this paper selected wood anatomical features from families of Magnoliidae and “lower” Hamamelididae are summarized and mapped onto previously published cladograms as a preliminary means of testing previous hypotheses of wood evolution. The results show that many of the characters are homoplasious and have evolved both in accord with, and counter to, the hypothesized general trends in different groups of flowering plants. In general, changes that confirm generalized trends are more common than changes that are counter to those trends. Future studies should combine wood anatomical characters with other features as part of a cladistic analysis. Fossil woods have not yet contributed significantly to phylogenetic studies, but in the very few cases where they have been linked to fossil reproductive structures, the woods have provided a better understanding of wood anatomy in early members of some families. Data from fossil wood expand the diversity of anatomical structure known in some angiosperm taxa and thus provide additional evidence that might be used in phylogenetic analyses. Fossil woods have the greatest potential to affect phylogenetic analyses where they can be linked to other fossil organs. The best chance for establishing such a linkage is through the study of fossil charcoalified woods that co-occur with other dispersed mesofossils.     

DEVELOPMENTALLY VARIABLE,POLYMORPHIC WOODS IN CACTI

Sixteen genera of cacti were discovered to have polymorphic wood, that is, the plants produce one type of wood while young but a different type when older. The polymorphisms are: fibrous wood (with vessels and scanty paratracheal parenchyma) followed by parenchymatous wood (with vessels but few or no fibers) (Hylocereus venezuelensis, Dendrocereus nudiflorus, Borzicactus humboldtii, Haageocereus australis, Morawetzia sericata, Stephanocereus leucostele, Trichocereus schickendantzii); WBT wood (with wide-band tracheids, vessels, and apotracheal parenchyma but few or no fibers) followed by fibrous wood (Buiningia aurea, Oreocereus celsianus, Vatricania guentheri); WBT wood followed by parenchymatous wood (Echinopsis tubiflora, Gymnocalycium marsoneri, G. oenanthemum, Notocactus warasii, Parodia maassii); trimorphic wood in which WBT wood is followed by fibrous wood, which is followed by parenchymatous wood (Melocactus intortus, Arrojadoa braunii). The different phases within each plant may differ in vessel cluster size, percentage of the vessels that are solitary, diameter of vessels, and lignification of ray cells. Several of these genera are not closely related to the others, so wood polymorphism may have arisen several times.     

Presence of paratracheal water storage tissue does not alter vessel characters in cactus wood

This research tested hypotheses that the presence of water storage tissues immediately adjacent to vessels would protect vessels from cavitation and would result in evolution of broader vessels that occur in fewer, smaller clusters relative to vessels surrounded by a matrix of fibers. We examined 21 species that have dimorphic wood, that is, at one stage in their life they produce a wood with a fibrous matrix surrounding the vessels and at another stage they produce wood with abundant paratracheal parenchyma or wide-band tracheids. In only one species were vessels in the water storage matrix broader than those in the fibrous matrix of the same plant. In most specimens, fibrous wood had smaller clusters of vessels than water storage wood, and a greater percentage of vessels in fibrous wood were solitary. Presence of abundant paratracheal water storage tissue was not correlated with a reduced number or size of rays. Axial masses in fibrous wood were not consistently narrower than those of water storage wood, consequently their vessels were not consistently closer to water stored in rays. Wood strength may be more important than conduction safety in determining vessel cluster size and widths of rays and axial masses.     

Vessel grouping patterns in subfamilies Apocynoideae and Periplocoideae confirm phylogenetic value of wood structure within Apocynaceae

This study contributes to our understanding of the phylogenetic significance and major evolutionary trends in the wood of the dogbane family (Apocynaceae), one of the largest and economically most important angiosperm families. Based on LM and SEM observations of 56 Apocynoideae species-representing all currently recognized tribes-and eight Periplocoideae, we found striking differences in vessel grouping patterns (radial multiples vs. large clusters) between the mainly nonclimbing apocynoid tribes (Wrightieae, Malouetieae, Nerieae) and the climbing lineages (remaining Apocynoideae and Periplocoideae). The presence of large vessel clusters in combination with fibers in the ground tissue characterizing the climbing Apocynoideae and Periplocoideae clearly contrasts with the climbing anatomy of the rauvolfioids (solitary vessels plus tracheids in ground tissue), supporting the view that (1) the climbing habit has evolved more than once in Apocynaceae, (2) the three nonclimbing apocynoid tribes are basal compared to the climbing apocynoids, and (3) Periplocoideae belong to the crown clade. The wood anatomy within the nonclimbing and climbing lineages is rather homogeneous, although a combination of specific characters (e.g. presence of septate fibers, axial parenchyma distribution, abundance of uniseriate compared to multiseriate rays, and presence and location of prismatic crystals) may be used to identify several tribes.     

The role of wood anatomy in phylogeny reconstruction of Ericales

The systematic significance of wood anatomical characters within Ericales is evaluated using separate and combined parsimony analyses including 23 wood characters and 3945 informative molecular characters. Analyses of wood features alone result in poorly resolved and conflicting topologies. However, when pedomorphic character states are coded as inapplicable, the combined bootstrap topology results in an increase of resolution and support at most deeper nodes compared with the molecular analyses. This suggests that phylogenetic information from the limited number of morphological characters is not completely swamped by an overwhelming amount of molecular data. Based on the morphology of vessels and fibers, and the distribution of axial parenchyma, two major wood types can be distinguished within Ericales: (i) a “primitive” type, nearly identical to the wood structure in the more basal outgroup Cornales, which is likely to have persisted in one major clade, and (ii) a “derived” type that must have evolved in at least two separate evolutionary lines. The occurrence of the first type is strongly correlated with shrubs to small trees growing in cold temperate or tropical montane regions, while the second type is common in tall trees of tropical lowlands. This favors the inclusion of ecologically adaptive features in phylogeny reconstruction. © The Willi Hennig Society 2006.     

Variation in the relationship between numbers of breeding pairs and woodland area for passerines in fragmented habitat

Species may differ in the relationship between the numbers of breeding pairs present and woodland area, because the proportion of a wood that forms suitable habitat will vary with woodland size. In this paper, we examine the pattern of variation in abundance with woodland area for eight breeding bird species, and also show how this pattern varied between years. During 1990-1997, we made annual censuses of 53–160 woods, of up to 10 ha in size, and fitted a power function to describe the relationships between numbers of breeding pairs and woodland area. Seven of the eight species, blackbird Turdus merula , dunnock Prunella modularis , wren Troglodytes troglodytes , great tit Parus major , chaffinch Fringilla coelebs , robin Erithacus rubecula and blue tit Parus caeruleus showed a pattern of proportionally higher numbers in smaller woods. Only long-tailed tit Aegithalos caudatus occurred in proportionally higher numbers in larger woods. Blackbird and dunnock showed a trend towards lower numbers in large woods during years with low regional population levels; for these species large woods may provide sub-optimal habitat. Great tit, blue tit, chaffinch and robin showed the opposite trend, towards lower numbers in small woods during years with low regional population levels; for these species small woods may provide sub-optimal habitat. Wren and long-tailed tit, which also showed large annual population fluctuations, showed no change in distribution with regional population level. In great tit and chaffinch, the distribution of pairs in any one year may have been influenced by site fidelity producing a lag in the response associated with regional population levels.     

Convergent evolution in discomycetes from bark and wood

Dry weathered wood (lignum and arid bark support a discomycete flora which is taxonomically extremely diverse and contains a high percentage of species with characters not found in Discomycetes of more mesic habitats. A large proportion of filiform-spored, and all muriform-spored non-lichenized Discomycetes occur in this habitat. Most lignicolous Discomycetes have either a pigmented epithecium or a dark margin which covers the hymenium when dry and many have reviving, gelaious ascomata. Their asci often have a simplified structure difficult to interpret. Weathered wood is a niche which has long continuity in geologic time but is relatively unfavorable for fungi because of exposure, aridity and low nutrient levels. Strong selective pressure has consequently been exerted on those fungi able to colonize it, resulting in a high incidence of convergent evolution.     

Wood anatomy and wood density in shrubs: Responses to varying aridity along transcontinental transects

Wood density plays a key role in ecological strategies and life history variation in woody plants, but little is known about its anatomical basis in shrubs. We quantified the relationships between wood density, anatomy, and climate in 61 shrub species from eight field sites along latitudinal belts between 31° and 35° in North and South America. Measurements included cell dimensions, transverse areas of each xylem cell type and percentage contact between different cell types and vessels. Wood density was more significantly correlated with precipitation and aridity than with temperature. High wood density was achieved through reductions in cell size and increases in the proportion of wall relative to lumen. Wood density was independent of vessel traits, suggesting that this trait does not impose conduction limitations in shrubs. The proportion of fibers in direct contact with vessels decreased with and was independent of wood density, indicating that the number of fiber-vessel contacts does not explain the previously observed correlation between wood density and implosion resistance. Axial and radial parenchyma each had a significant but opposite association with wood density. Fiber size and wall thickness link wood density, life history, and ecological strategies by controlling the proportion of carbon invested per unit stem volume.     

Palaeoecology of the Miocene El Cien Formation (Mexico) as determined from wood anatomical characters

In order to establish the general environmental conditions under which the plants of the El Cien Formation grew during the Miocene, wood anatomical features from several extant ecological categories (southern California flora) and communities (Brazil, Mexico and Suriname) as well as those of the El Cien paleocommunity were analysed using different statistical approaches. We find a significant correlation between 42 characters of 199 species and 11 climate variables which suggests a close association of wood characters with temperature, whereas characters such as diameter, frequency, and grouping of vessels are more closely related to evaporation. The expression of various wood characters depends on more than one climate variable. There are no doubts regarding the tropical affiliation of the El Cien fossil flora but water availability and seasonality of the system are less clear. Principal component and cluster analyses of two different datasets (423 species, 9 characters and 223 species, 32 characters) suggest that the physiognomy of the fossil assemblage of El Cien was highly efficient in water transport, but also highly vulnerable to embolisms. Characters such as vessel diameter, frequency and grouping suggest proximity to a tropical rain forest, whereas the proportion of ring semi ring porous woods brings the El Cien paleoflora closer to forests with slightly xeromorphic physiognomy, probably a tropical deciduous or semi-deciduous forest. This condition is paralleled with the taxonomic composition of the El Cien fossil plants since their Nearest Living Relatives (NLR) can be referred to both humid or more (seasonally) dry environments. Derived regression analysis models suggest that the paleoflora prospered under conditions with an evaporation regime similar to that of the tropical rain forest.     

Wide-band tracheids are present in almost all species of Cactaceae

Wide-band tracheids (WBTs) have been found in seedlings of most species of cacti that have fibrous wood in their adult bodies. Consequently, this cell type is now known to be present in almost all cacti. Earlier studies of adult plants revealed WBTs to be present only in cacti with globose or short, broad bodies, whereas all species with large columnar or long slender bodies had fibrous wood without WBTs. However, even these species produce WBTs during the first several months after germination. In species with fibrous wood in their adult bodies (species with large or slender bodies), seedlings undergo a phase transition in wood morphogenesis after a few months and stop producing the juvenile (WBT) wood and begin producing adult (fibrous) wood. If adult plants have an intermediate size, the phase transition is delayed and the plant produces WBT wood for several years. Species with globose bodies repress the phase transition completely and never switch to producing adult (fibrous) wood. Because WBTs are so widespread, they probably originated only once in Cactaceae, not multiple times as suggested earlier, or there may have been just a single origin in the Cactaceae/Portulacaceae clade.     

Dicotyledonous woods from the Upper Cretaceous of southern Illinois

Dicotyledonous woods from the Upper Cretaceous of Southern IllinoiS. Five species of fossil dicotyledonous wood are described from an Upper Cretaceous (Maestrichtian; locality in Alexander County, IllinoiS. U.S. A. Paraquercimum cretaceum has structure similar to the Fagaceae (evergreen Oak- Lithocarpus ) and Casuarinaceae and represents the earliest known occurrence of this structural type (large solitary pores and uniseriate and large multiseriale rays). Paraphyltanthoxyhin illirioisense and Icacinoxylon alternipunctata are species of genera represented at other Cretaceous and Early Tertiary localities In large diameter trees. Parabombacaceoxylon magniporosum has large diameter pores and scalariform perforation plates, a combination of characters that is extremely rare in the extant flora. Paraapocynaceoxylon barghoorni has a combination of characters represented in extant Apocynaceae. These five species lack growth rings, have high vulnerability indices (mean vessel diameter divided by mean number of vessels per square millimeter, and a relatively high proportion of ray parenchyma. They lack specialized wood anatomical characters, and a compilation of vessel element lengths in these and other Cretaceous woods indicates that short vessel elements (a derived character) were less frequent in the Cretaceous than in extant dicotyledonous trees.     

Ontogenetic mechanisms and the evolution of Cactaceae

Abstract

The ancestors of cacti were leafy trees that had hard, woody trunks. The development of the cactus body is controlled by ontogenetic mechanisms that have evolved, and now they produce a body that is leafless, succulent and has a photosynthetic cortex. Specific changes include: bark formation is postponed and the epidermis and stomata function for many years; the outer cortex is a palisade cortex with intercellular spaces; there are cortical bundles that resemble leaf veins but which have secondary xylem and phloem. Wood development has changed dramatically such that water storage is maximized (increased ray parenchyma) and danger of water stress is minimized (increased paratracheal parenchyma, loss of fibers). Several genera have polymorphic wood: the plants produce one type of wood for several years, then later they produce a different type. It is possible that the extensive evolutionary changes have resulted from mutations in the controller regions of genes, not in the structural regions.     

Factors influencing Willow Tit Poecile montanus site occupancy: a comparison of abandoned and occupied woods

The British Willow Tit Poecile montanus kleinschmidti underwent a decline of 85% between 1970 and 2003. The cause of this decline is unknown. However, several hypotheses have been put forward to account for it: competition from other tit species, predation by Great Spotted Woodpeckers Dendrocopos major and habitat change. In order to test these, woods that are currently occupied by Willow Tits were paired with woods (within 50 km) that had been abandoned by Willow Tits five or more years previously. Point counts for other tit species (potential competitors) and woodpecker species (potential predators) were carried out at ten evenly spaced points throughout each wood. Habitat variables were collected within a 50-m radius of where a Willow Tit was located (in the occupied woods) or where maps showed a Willow Tit had been located (for abandoned woods). No evidence was found for differences in numbers of potential competitor or potential predator species in abandoned and occupied woods. Soil water content was found to be higher at occupied sites. No other habitat features differed between the two categories of site. The drying up of British woods could therefore be implicated in the Willow Tit decline and this warrants further investigation.     

Universal hydraulics of the flowering plants: vessel diameter scales with stem length across angiosperm lineages,habits and climates

Angiosperm hydraulic performance is crucially affected by the diameters of vessels, the water conducting conduits in the wood. Hydraulic optimality models suggest that vessels should widen predictably from stem tip to base, buffering hydrodynamic resistance accruing as stems, and therefore conductive path, increase in length. Data from 257 species (609 samples) show that vessels widen as predicted with distance from the stem apex across angiosperm orders, habits and habitats. Standardising for stem length, vessels are only slightly wider in warm/moist climates and in lianas, showing that, rather than climate or habit, plant size is by far the main driver of global variation in mean vessel diameter. Terminal twig vessels become wider as plant height increases, while vessel density decreases slightly less than expected tip to base. These patterns lead to testable predictions regarding evolutionary strategies allowing plants to minimise carbon costs per unit leaf area even as height increases.     

WOOD ANATOMY OF THE AMERICAN FRANKENIAS (FRANKENIACEAE): SYSTEMATIC AND EVOLUTIONARY IMPLICATIONS

Results of a comparative wood anatomical survey of the American frankenias are presented. The eleven species examined are woody perennials occurring almost exclusively in arid and semiarid regions and on saline and gypseous soils. The secondary xylem of all species is highly specialized and is characterized by libriform fibers, vessel elements with simple perforation plates, and the absence of rays. Axial elements of all species are quite small. A number of unusual features, e.g., anomalous secondary growth and formation of interxylary cork, were observed in some species. Nonfibrous woods have evolved independently in two species of reduced stature and contrast markedly with the highly fibrous woods of most species. Woods of the American frankenias are compared with those of the Tamaricaceae. The systematic and evolutionary implications of interspecific variation in both qualitative and quantitative features are discussed. There is a general tendency for dimensions of the axial wood elements to be positively associated and to decrease with decreasing plant height. In general, differences in wood anatomy more strongly reflect differences in plant growth form and size than phylogeny.