Differences in patterns of cell death between ray parenchyma cells and ray tracheids in the conifers <Emphasis Type="Italic">Pinus densiflora</Emphasis> and <Emphasis Type="Italic">Pinus rigida</Emphasis>

Differences in patterns of cell death between ray parenchyma cells and ray tracheids in the conifers Pinus densiflora and Pinus rigida were clarified. Differentiation and cell death of ray tracheids occurred successively and both were related to the distance from the cambium. In this respect, they resembled those of longitudinal tracheids. Thus, the cell death of short-lived ray tracheids could be characterized as time-dependent programmed cell death. In contrast, ray parenchyma cells survived for several years or more, and no successive cell death occurred, even within a single radial line of cells in a ray. Thus, the features of death of the ray parenchyma cells were different from those of ray tracheids. Cell death occurred early in ray parenchyma cells that were in contact with ray tracheids. The initiation of secondary wall thickening occurred earlier in ray parenchyma cells that were in contact with ray tracheids in Pinus densiflora than in others. In addition, localized thickening of secondary walls occurred only in ray parenchyma cells that were in contact with ray tracheids in Pinus rigida. Moreover, no polyphenols were evident in such cells in either species. Therefore, ray parenchyma cells that were in contact with ray tracheids appeared not to play a role in the formation of heartwood extractives. Our observations indicate that short-lived ray tracheids might affect the pattern of differentiation and, thus, the functions of neighboring long-lived ray parenchyma cells in conifers.     

Pinoid woods with resin canals from the upper cretaceous of Hokkaido and Saghalien

This paper describes six species of permineralized pinoid woods with resin canals from the Upper Cretaceous of Hokkaido, Japan and Saghalien:Piceoxylon transiens Shimakura,P. scleromedullosum Shimakura,P. macroporosum sp. nov.,P. takahashii sp. nov.,Pinuxylon microporosum Ogura andPinoxylon dakotense (Knowlton) Read (new to Saghalien and the Cretaceous).Piceoxylon macroporosum is characterized by large resin canals and rays without ray tracheids and in lacking tylosoids in resin canals.Piceoxylon takahashii, which resemblesP. scleromedullosum, is distinguished from the latter in having ray tracheids and nests of sclereids in pith. Structure and affinities of the petrified plants from the Cretaceous of Northern Japan and Saghalien XVI. (continued from Ohsawaet al. 1995)     

Nuclear DNA fragmentation during cell death of short-lived ray tracheids in the conifer <Emphasis Type="Italic">Pinus densiflora</Emphasis>

One key event in the programmed cell death is nuclear DNA fragmentation. We investigated the timing of nuclear DNA fragmentation during the cell death of short-lived ray tracheids in Pinus densiflora using the terminal deoxynucleotidyl transferase-mediated dUTP nick-end labeling (TUNEL) assay. Fluorescence due to TUNEL was detected only in deformed nuclei that lacked obvious chromatin in ray tracheids that were adjacent to ray tracheids that no longer contained nuclei. Our observations revealed that nuclear DNA fragmentation occurred only at the final stage of cell death in ray tracheids in situ.     

SILICIFIED PINUS REMAINS FROM THE MIOCENE OF WASHINGTON

Silicified leaves, dwarf shoots, pollen cones, and seed cones of Pinus from a Late Miocene chert bed within the Yakima Basalt Formation near Yakima, Washington are interpreted as coming from a single new species, P. foisyi. The needles and dwarf shoots are those of a three-needle pine. The needles contain two to four medial resin canals, a biform hypodermis, and endodermal cells with uniformly thickened walls. The pollen cones are ellipsoidal and about 1 cm long, and many contain bisaccate pollen grains. The seed cones are at least 6 cm long and are slightly asymmetrical. The cone axis has a broad sclerotic outer cortex, and the seed wing extends from a thick parenchymatous base. The scale apex bears a conspicuously swollen projection. The foliage and seed cones are identifiable with the Subgenus Pinus, Section Pinus, Subsection Oocarpae independently of one another, and together indicate a fossil species related to the modem Californian closed cone pines. Pinus foisyi represents one of the earliest occurrences of cone asymmetry associated with this group. However, cone serotiny characteristic of the modem species appears to have evolved after the Late Miocene.     

WOOD ANATOMY OF BUBBIA (WINTERACEAE), WITH COMMENTS ON ORIGIN OF VESSELS IN DICOTYLEDONS

Quantitative and qualitative features of wood anatomy are reported for ten collections of seven species of Bubbia. Variations on the basic plan for Winteraceae can be interpreted in terms of taxonomic and ecological distinctions. Tracheid length is correlated with plant size and habit: tracheids are shortest in shrubs. Tracheid wall thickness and ray cell wall thickness distinguish species. Ray cell procumbency and multiseriate ray width increase with age. Growth rings occur only in a species from stream margins. SEM studies reveal absence of a warty layer within tracheids. Helical thickenings are absent. Presence of these two features in Pseudowintera may be correlated with the cool temperate habitats of that genus. Overlap areas of tracheids in Bubbia show various degrees of scalariform pitting, ranging from none (B. semecarpoides) to abundant presence (B. balansae). Perforation-like pits in tracheids of the latter prove, with SEM studies, to have pit membranes containing porosities less than 1 μm in diameter. Scalariform pitting on overlap areas is absent in earlier secondary xylem and increases during later secondary xylem. Scalariform lateral wall pitting can occur in abnormally wide tracheids formed after pauses in cambial activity. These facts show that primitive dicotyledon woods like those of Bubbia can activate genetic information for scalariform end wall patterns and lateral wall pitting such as primitive vessels show without the intervention of paedomorphosis. Paedomorphosis in dicotyledon woods is held still to apply only to special herbaceous and herblike growth forms, not to primarily woody plants. Progenesis (in xylem, loss of secondary xylem) is not held to be necessary to account for the scalariform patterns seen in tracheary elements of primitive dicotyledons. Reasons are given for rejection of the hypothesis that Winteraceae and other woody dicotyledons (Amborella, Sarcandra, Tetracentron, Trochodendron) are secondarily vesselless.     

Nucleotide diversity and demographic history of Pinus bungeana,an endangered conifer species endemic in China

Orographic and climatic oscillations have played crucial roles in shaping the nucleotide diversity and evolutionary history of many species across the Northern Hemisphere. In this study, based on 10 nuclear loci and a chloroplast DNA marker, we analyzed the nucleotide polymorphisms and demographic history of the endangered conifer species Pinus bungeana in Northwest China and investigated the phylogenetic relationships between P. bungeana and two related species, that is, Pinus gerardiana and Pinus squamata. We found that P. bungeana exhibited an extremely low level of nucleotide diversity (π_sil = 0.00159). Demographic simulations based on DIYABC analysis showed that P. bungeana underwent demographic expansion and contraction during the Miocene. According to ecological niche modeling, we found that this species survived in situ during the glacial period and was not restricted to southern refugia. We speculate that P. bungeana may have experienced widespread population shrinkage from the Last Interglacial to the Last Glacial Maximum due to geological or climatic events. Isolation‐with‐migration analysis revealed that the divergence (~2.4–4.2 Ma) among P. bungeana and its related species was significantly associated with the Qinghai–Tibetan Plateau uplift events in the mid‐to‐late Tertiary period. Species tree analyses suggested that these three related Pinus species formed a monophyletic clade with high bootstrap support. These results suggest that the Miocene–Pliocene and Pleistocene geological and climatic fluctuations might have profoundly affected the nucleotide diversity and demography of this psychrotolerant conifer species in western China.     

Structural analysis of K<Superscript>+</Superscript> dependence in <Emphasis Type="SmallCaps">l</Emphasis>-asparaginases from <Emphasis Type="Italic">Lotus japonicus</Emphasis>

Wood is composed of various types of cells and each type of cell has different structural and functional properties. However, the temporal and spatial diversities of cell wall components in the cell wall between different cell types are rarely understood. To extend our understanding of distributional diversities of cell wall components among cells, we investigated the immunolabeling of mannans (O-acetyl-galactoglucomannans, GGMs) and xylans (arabino-4-O-methylglucuronoxylans, AGXs) in ray cells and pits. The labeling of GGMs and AGXs was temporally different in ray cells. GGM labeling began to be detected in ray cells at early stages of S₁ formation in tracheids, whereas AGX labeling began to be detected in ray cells at the S₂ formation stage in tracheids. The occurrence of GGM and AGX labeling in ray cells was also temporally different from that of tracheids. AGX labeling began to be detected much later in ray cells than in tracheids. GGM labeling also began to be detected in ray cells either slightly earlier or later than in tracheids. In pits, GGM labeling was detected in bordered and cross-field pit membranes at early stages of pit formation, but not observed in mature pits, indicating that enzymes capable of GGM degradation may be involved in pit membrane formation. In contrast to GGMs, AGXs were not detected in pit membranes during the entire developmental process of bordered and cross-field pits. AGXs showed structural and depositional variations in pit borders depending on the developmental stage of bordered and cross-field pits.     

CONIFER WOOD FROM THE UPPER JURASSIC OF UTAH PART I: XENOXYLON MORRISONENSE SP. NOV.

A new species of conifer wood, Xenoxylon morrisonense, is described from the Morrison Formation on the Colorado Plateau. It is compared with other species of Xenoxylon, with X. latiporosum being the closest. Xenoxylon morrisonense differs from X. latiporosum in its marked indentations, simple pits on the horizontal and tangential walls of ray cells, absence of crassulae, presence of wood parenchyma, and thin borders on podocarpoid type crossfield pits. The origin of the septa in the tracheids is summarized, and the possible affinity of Xenoxylon with the Podocarpaceae is considered.     

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.     

INCREASE IN CELL WALL SURFACE AREA DURING ENLARGEMENT OF CAMBIAL DERIVATIVES IN ABIES CONCOLOR

Wilson , Brayton F. (U. California, Berkeley). Increase in cell wall surface area during enlargement of cambial derivatives in Abies eoncolor . Amer. Jour. Bot. 50(1): 95–102. Illus. 1963.— Dimensions of fusiform cells (tracheids and sieve cells) and ray cells were measured from samples of the 1960 xylem and phloem increment of 5 trees felled at monthly intervals from April through July, 1960. Calculations using these measurements gave the magnitude, direction and rate of increase in cell wall surface area during enlargement. Although 14 times more tracheids than sieve cells were produced, both cell types enlarged mostly in a radial direction (up to 400%) at the same rate (20–33 × 10³μ² wall surface area/day) to the same final size. Fusiform cambial cells doubled their wall area between successive periclinal divisions. Calculations showed that ⅞ of this increase was in the radial walls of the daughter cells at a rate comparable to that in enlarging tracheids and sieve cells; the other ⅞ was from cell plate formation at an estimated rate of 187–327 × 10³μ²/day. Enlargement of derivatives in the radial direction largely determined the amount of increase in wall area. Besides radial enlargement, tracheids also elongated (up to 13%) and phloem cells enlarged tangentially (sieve cells up to 36%; pholem ray cells up to 60%). The relationships of enlarging tracheids and xylem ray cells are discussed, and it is suggested that slippage may occur between the developing walls.     

EQUISETUM CLARNOI,A NEW SPECIES BASED ON PETRIFACTIONS FROM THE EOCENE OF OREGON

Equisetum clarnoi is described from four silicified stem fragments and numerous small roots from the Eocene Clarno Chert of Jefferson County, Oregon. Stems are up to 8.0 mm in diam and have sunken stomata arranged vertically in a single line flanking each of the external biangulate stem ridges, features that clearly ally this species with the subgenus Hippochaete. External stem ridges are equal in number to the carinal hypodermal bands. The hypodermis is composed of fibers and has prominent carinal bands up to 0.75 mm long and shorter vallecular bands. Cortical parenchyma cells enclose prominent vallecular canals which are lined by specialized thick-walled parenchyma cells. The double, common endodermis has prominent casparian strips. Vascular bundles are composed of four to seven metaxylem tracheids flanking each side of the phloem and protoxylem tracheids which occur singly on the internal surface of the small carinal canals. Leaf sheaths in cross section have an adaxial fibrous layer and an external or near external fibrous bundle. Roots are up to 2.0 mm in diam and have paired cuboidal epidermal cells from which root hairs arise. The stele of the root is central and shows exarch primary xylem maturation. Equisetum clarnoi most closely resembles the extant Equisetum hyemale var. affine.     

Freezing behavior of xylem ray parenchyma cells in softwood species with differences in the organization of cell walls

Summary By cryo-scanning electron microscopy we examined the effects of the organization of the cell walls of xylem ray parenchyma cells on freezing behavior, namely, the capacity for supercooling and extracellular freezing, in various softwood species. Distinct differences in organization of the cell wall were associated with differences in freezing behavior. Xylem ray parenchyma cells with thin, unlignified primary walls in the entire region (all cells inSciadopitys verticillata and immature cells inPinus densiflora) or in most of the region (mature cells inP. densiflora and all cells inP. pariflora var.pentaphylla) responded to freezing conditions by extracellular freezing, whereas xylem ray parenchyma cells with thick, lignified primary walls (all cells inCrytomeria japonica) or secondary walls (all cells inLarix leptolepis) in most regions responded to freezing by supercooling. The freezing behavior of xylem ray parenchyma cells inL. leptolepis changed seasonally from supercooling in summer to extracellular freezing in winter, even though no detectable changes in the organization of cell walls were apparent. These results in the examined softwood species indicate that freezing behavior of xylem ray parenchyma cells changes in parallel not only with clear differences in the organization of cell walls but also with subtle sub-electron-microscopic differences, probably, in the structure of the cell wall.     

PERMINERALIZED PINACEOUS LEAVES FROM THE UPPER CRETACEOUS OF HOKKAIDO

Three fragments of fascicles of a possible five-needle pine are described from the Upper Cretaceous Yezo Group (Santonian/Senonian) of Hokkaido. Specimens from the Omakizawa, Oyubari, Yubari City, the Koyanozawa, Ikushumbetsu, Mikasa City, and the Sankebetsugawa, Haboro are preserved in calcium carbonate nodules containing abundant ammonites. Leaves borne in apparent fascicles of five measure 0.7–0.8 mm in radial and 0.8–1.0 mm in tangential diam and are represented by short fragments of isolated needles. Thick-walled epidermal cells on these amphistomatic leaves resemble the underlying uniform hypodermis. Two external resin canals are situated near the lateral corners toward the adaxial surface. Only slightly plicate mesophyll cells in a layer one to two cells thick border on an elliptical endodermis with thickened outer cell walls. Two to three layers of transfusion tissue surround the double vascular strand. Two vascular bundles are separated by one cell layer of sclerenchyma fibers. Small patches of abaxial and adaxial sclerenchyma fibers have also been observed. Leaves most closely resemble those of Pinus leiophylla Schl. et Cham. Subgenus Pinus, Section Pinea, Subsection Leiophyllae and P. montezumae Lamb. Subgenus Pinus, Section Pinus, Subsection Ponderosae and are described as a new species P. hokkaidoensis sp. nov. Stockey and Ueda. Close anatomical comparisons are made with this leaf and previously described permineralized Upper Cretaceous pine needles from Hokkaido and eastern North America.     

Variation in wood anatomical traits of Pinus sylvestris L. between Spanish regions of provenance

The Spanish populations of Pinus sylvestris L. occupy differentiated sites and must therefore include structural variations to cope with varied climate conditions. This study compares wood anatomical traits of P. sylvestris from ten Spanish regions of provenance with contrasting climates, taking into account the effects of region of provenance and tree nested within provenance on variation in wood biometry. In general, the effect of both sources of variation (provenance and tree) on wood biometry was highly significant. Most of the anatomical variations observed were intra-populational (at the tree level), although variation explained by provenance was high for some parameters (e.g., ray frequency and ray parenchyma cell frequency), suggesting high environmental influence. Trees in the driest region, growing in a Mediterranean phytoclimate, were characterized by large tracheid lumens, suggesting more efficient water conduction. They also had thick cell walls, which would reduce the risk of cavitation caused by high implosion stress during periods of drought, as well as a high ray tracheid frequency, implying greater water storage capacity in the sapwood. The population with greatest growth, located in an oroboreal phytoclimate, was characterized by large bordered pits and long tracheids, which would reduce resistivity in water transport. At higher altitudes, tracheid lumen diameter and resin canal diameter tended to be smaller, and intertracheid wall strength was greater. Results are discussed in relation to adaptation of the species to growth demands and frost.     

A new fossil Cedrus species from the early Miocene of northwestern Turkey and its possible affinities

A new fossil species, Cedrus anatolica n. sp., is described from the early Miocene Hançili Formation of Turkey. All analyses were performed on the thin sections housed at Istanbul University – Cerrahpasa. The new species was interpreted as having the closest affinity with the modern Mediterranean species Cedrus atlantica (Endlicher) Manetti ex Carriere and Cedrus libani Richard. The evolutionary line shows some changes in wood anatomy. From the early Cretaceous to the early Miocene, the pits on the tangential walls of the tracheids gradually decreased, the height of rays increased and the number of epithelial cells in the traumatic resin canals increased slightly. These features are similar in three modern species; other wood anatomical features are also quite stable among the new fossil and modern species.     

Discovery of fossil Ayu (<Emphasis Type="Italic">Plecoglossus altivelis</Emphasis>) from the Upper Miocene of Shimane Prefecture,Japan

The Ayu, Plecoglossus altivelis, comprising three subspecies, is the only species in the genus Plecoglossus in the family Osmeridae. Here, we describe for the first time the discovery of fossil specimens of P. altivelis from the Upper Miocene sediment of Matsue, Shimane Prefecture, Japan. The fossil individuals are subadult and have comb teeth in the second and third generations. These fossil specimens are clearly similar to the recent P. altivelis, the former’s meristic characters being included in the three recent subspecies. In terms of numbers of pectoral fin rays and vertebrae, the fossil specimens are more similar to the subspecies Plecoglossus altivelis ryukyuensis than the other subspecies, but they also show different dorsal fin shapes and anal fin ray counts. This discovery of fossil P. altivelis indicates the persistence of the species for at least 10 million years, from the Late Miocene to the present.     

A FURTHER CONTRIBUTION TO CLOSED-CONE PINE (OOCARPAE) HISTORY

A new record of a fossil Monterey pine (Pinus radiata D. Don) is recorded from rocks of the Plio-Pleistocene age, southern California, and the affinities of other presumed fossil closed-cone pines are revised. This has led to the recognition of two new pine species, Pinus storeyana Axelrod of subsect. Oocarpae and Pinus verdiana Axelrod of subsect. Ponderosae. Geologic evidence indicates that forerunners of the present California and Mexican species of Oocarpae were in proximity in Mexico during the Miocene. They were isolated following opening of the Gulf of California, the northward movement of Baja and Alta California on the San Andreas fault system, and the spread of desert climate. New populations of Pinus radiata developed during the Pliocene (5–2 million years) and later as environmental diversity increased. This also enabled Pinus remorala Mason and P. muricata D. Don, which the fossil record suggests were distinct into the Late Pleistocene, to hybridize in the new topographic-climatic Holocene environments.     

STRUCTURE OF TRACHEIDS IN THREE SPECIES OF LYCOPODIUM

The structure of tracheids in Lycopodium lucidulum, L. clavatum, and L. tristachyum was studied with the light microscope. Protoxylem development is at least sometimes and possibly always mesarch in indeterminate axes of all three species. Centrifugally formed protoxylem elements are reticulate and discontinuities in the secondary walls of these elements are sometimes conspicuously bordered. Wall thickenings of first formed protoxylem elements consist mainly of indirectly connected rings. Late centripetally formed protoxylem elements and transitional elements have a reticulate secondary wall pattern. The narrowest metaxylem elements have circular bordered pits while in wider metaxylem elements pits are bordered and may vary from circular to scalariform. Pitting is uniseriate to triseriate in tracheids of all three species, and intermittent tetraseriate pitting was occasionally observed in L. lucidulum. Crassulae occur in tracheids of the three species, and in L. clavatum an additional framework, probably representing thickened compound middle lamella, is also present. Pits often appear helically arranged, and in all three species pits are connected by thin areas in the secondary wall. Macrofibrils approximately 0.5 μ wide were observed in tracheids of the three species. In L. clavatum the arrangement of macro-fibrils was predominantly bidirectional.     

The positional distribution of cell death of ray parenchyma in a conifer, Abies sachalinensis

We monitored the distribution of death of secondary xylem cells in a conifer, Abies sachalinensis. The cell death of tracheids, which are tracheary elements, occurred successively and was related to the distance from cambium. Thus, it resembled programmed cell death. By contrast, the death of long-lived ray parenchyma cells had the following features: (1) ray parenchyma cells remained alive for several years or more; (2) in many cases, no successive cell death occurred even within a given radial cell line of a ray; and (3) the timing of cell death differed among upper and lower radial cell lines and other lines of cells within a ray. These results indicate that the death of long-lived ray parenchyma cells involves a different process from the death of tracheids. The initiation of secondary wall formation and the lignification of ray parenchyma cells in the current year's annual ring were delayed in the upper and lower radial cell lines of a ray. In addition, the density of distribution and orientation of cortical microtubules in such cells were different from those in cells in other radial lines. Ray parenchyma cells in the previous year's annual ring within the upper and lower radial cell lines of a ray contained many starch grains. Our results indicate that positional information is an important factor in the control of the pattern of differentiation and, thus, of the functions of ray parenchyma cells that are derived from the same cambial ray cells.     

The anatomy of the chi-chi of <Emphasis Type="Italic">Ginkgo biloba</Emphasis> suggests a mode of elongation growth that is an alternative to growth driven by an apical meristem

The chi-chi of Ginkgo biloba L. are cylindrical woody structures that grow downwards from the branches and trunks of old trees, eventually entering the soil where they give rise to adventitious shoots and roots. Examination of segments of young chi-chi taken from a mature ginkgo tree revealed an internal woody portion with irregular growth rings of tracheid-containing secondary xylem covered by a vascular cambium and bark. The cambium was composed of both fusiform cells and parenchymatous ray cells. Near the tip of the chi-chi, these two types of cambial cells had orientations ranging between axial, radial and circumferential with respect to the cylindrical form of the chi-chi. The xylem rays and tracheids that derived from the cambium showed correspondingly variable orientations. Towards the base of the chi-chi, the fusiform cells and young tracheids were aligned parallel to the axis, indicating that the orientation of the cambial cells in basal regions of the chi-chi gradually became normalised as the tip of the chi-chi extended forwards. Nevertheless, in such basal sites, tracheids near the centre of the chi-chi showed variable orientations in accordance with their mode of formation during the early stages of chi-chi development. The initiation of a chi-chi is proposed to derive from a localised hyperactivity of vascular cambial-cell production in the supporting stem. The chi-chi elongates by tip growth, but it does so in a manner different from organ growth driven by an apical meristem. It is suggested that the chi-chi of Ginkgo is an “evolutionary experiment” that makes use of the vascular cambium, not only for its widening growth but also for its elongation.