Secondary Xylem Tracheids Research Articles

A new conifer stem, Yiwupitys elegans, from the Yiwu Jurassic Forest, Hami, Xinjiang, Northwest China

Fossil wood not only provides a comprehensive understanding of the ancient floral composition and the evolution history of wood through time, but also sheds light on the paleoclimate, paleoenvironment, and paleoecology. Permineralized conifer wood commonly occurs in the Jurassic of China; however, the detailed anatomical study has been sparsely conducted on the Jurassic wood from Xinjiang Uygur Autonomous Region (Xinjiang), Northwest China. Here, we describe a unique permineralized stem, Yiwupitys elegans Gou et Feng gen. nov. et sp. nov. from the Middle Jurassic Xishanyao Formation in Naomaohu Town of Yiwu County, Hami City, Xinjiang, Northwest China. The fossil stem is preserved with pith, primary xylem, and secondary xylem. The solid pith is heterocellular and characterized mainly by clustered or isolated secretory cells and pitted parenchymatous cells at the pith periphery. The primary xylem is mesarch showing annular, helical, and scalariform thickenings on the tracheid walls from the protoxylem to the metaxylem. The secondary xylem is pycnoxylic and exclusively composed of tracheids and parenchymatous rays. Growth rings are distinct. The secondary xylem tracheids are featured commonly by bent ends overlapping each other. They have mostly uniseriate or biseriate, or partially triseriate alternately or oppositely arranged bordered pits on the radial walls and uniseriate or biseriate bordered pits on the tangential walls. The rays are uniseriate and up to sixteen cells high. There are four to eight cupressoid pits in each cross-field, arranged in the araucarioid type. Leaf traces initiate from the pith margin and possess a single vascular strand in each trace. The anatomical structures indicate that Yiwupitys elegans has a closer affinity with conifers, although it shows few characteristics of ginkgophytes. Our study shed new light on the plant diversity of the Yiwu Jurassic Forest.

A new conifer stem, Ductoagathoxylon wangii from the Middle Jurassic of the Santanghu Basin, Xinjiang, Northwest China

A new permineralized stem, Ductoagathoxylon wangii Gou et Feng, is described from the Middle Jurassic Xishanyao Formation in Naomaohu Town of Yiwu County, Hami City, Xinjiang Uygur Autonomous Region, Northwest China. The fossil stem is preserved with pith, primary xylem, and secondary xylem. The solid pith is heterocellular, and characterized by the presence of regularly arranged clusters of secretory cells at the periphery. The primary xylem is endarch to mesarch, showing annular, helical, and scalariform thickenings on the tracheid walls from the protoxylem to the metaxylem. The secondary xylem is pycnoxylic and exclusively composed of tracheids and parenchymatous rays. The secondary xylem tracheids have mostly uniseriate, partially biseriate, and rarely triseriate alternately arranged contiguous bordered pits on their radial walls. The rays are uniseriate and up to ten cells high. There are five to nine cupressoid pits in each cross-field, arranged in the araucarioid type. Growth rings are distinct and are characterized by a narrow band of latewood. This is the third fossil stem described from the Middle Jurassic of China that is preserved with anatomical structures of pith, primary, and secondary xylem, thus shedding new light on the diversity and evolution of conifers. The occurrence of a traumatic scar in the stem indicates that the tree was probably damaged by an abiotic cause, but it successfully survived through physiological responses to the wound.

Severe insect defoliation at different timing affects cell wall formation of tracheids in secondary xylem of Larix kaempferi

Severe insect defoliation induced non-lignified tracheids and thin-walled latewood tracheids in the defoliation year in Larix kaempferi. These anomalous tracheids had heterogenous lignin deposition of S2 layer. Severe insect defoliation induces the formation of light ring consisted of thin-walled latewood tracheids in larch trees, which would affect wood quality. However, the identity of the secondary wall layers affected by insect defoliation remains unclear. The aim of this study is to clarify the effects of insect defoliation on the formation of secondary cell walls of tracheids in Larix kaempferi with focus on the defoliation timing. We observed by light microscopy and transmission electron microscopy (TEM) the secondary cell walls of tracheids produced in a defoliation year in L. kaempferi trees on which needles were attacked in July (gypsy moth, GM samples) or August (larch sawfly, LS samples). Light microscopy observations revealed that GM samples produced non-lignified tracheids in the transition zone between earlywood and latewood, as well as thin-walled latewood tracheids, and non-lignified tracheids were observed near the cambial zone in LS samples following defoliation for two consecutive years. TEM observations revealed that the tracheids in both samples had anomalous layers in secondary walls such as thin S2 layers with heterogeneous lignin deposition. The production of non-lignified tracheids corresponded to defoliation date in both samples, unlike changes in the layer composition of anomalous tracheids. Our study suggests that changes in wood structure depend on the date of insect defoliation and that insect defoliation affects the formation of secondary cell walls of tracheids, presumably in response to inadequate photosynthate supply as a result of defoliation.

Callixylon wendtiisp. nov., a new species of archaeopteridalean progymnosperm from the Late Devonian of Anti-Atlas, Morocco

ABSTRACTArchaeopterid trees were the main components of most Late Devonian forests. Their aerial axes characterised by a eustele with mesarch primary xylem strands, leaf traces departing radially from cauline bundles and secondary xylem tracheids with radial pits arranged in groups, are referred to the genusCallixylonZalessky. The nineteen species ofCallixylondescribed to date from North America, North Africa, Europe, Xinjiang and several parts of Russia range from the late Givetian to the Mississippian. In this paper, we describe a new species ofCallixylonfrom two specimens collected in the Famennian locality of Mader el Mrakib in eastern Anti-Atlas.Callixylon wendtiisp. nov. is characterised by the presence of sclerotic nests in the pith, a new character for the genus. Its wood shows narrow rays of variable height, with unevenly distributed ray tracheids. Small vascular traces crossing the wood within the two innermost growth rings are interpreted as evidence for short-lived leaves. This discovery adds to the diversity of the genusCallixylonin an area of northern Gondwana that may have been favourable to the establishment of a diverse community of archaeopterid trees.

Cell biology of bordered-pit formation in balsam-fir trees

A mature bordered pit in secondary xylem of Pinaceae comprises a circular border of secondary-wall material that protrudes into the tracheid lumen and is punctuated by a centralized aperture through which sap flows. The overarching border encloses a pit chamber within which is a “membrane”, or diaphragm, consisting of a central torus and margo strands. Bordered-pit pairs are abundantly present in all woods, and their membranes serve as swinging-diaphragm check valves regulating sap flow between adjoining tracheary elements, simultaneously trapping emboli and particulates in water as it moves from roots to leaves. The cell biology of bordered-pit formation in cambial derivatives of Abies balsamea (L.) Mill. was investigated by light and scanning electron microscopy during early stages of cellular differentiation of cambial derivatives into secondary xylem tracheids. A bordered-pit template (BPT), a bordered-pit organelle (BPO), a bordered-pit basal disk, and additional novel structures were found to be associated with bordered-pit formation. Evidence was found that the membrane does not comprise residual compound middle lamella; rather, the membrane forms de novo as BPO remnants. A cell-biology model and new terminology are introduced to explain how BPTs, BPOs, and basal disks contribute to successive stages in formation of bordered-pit pairs.

Apical control of xylem formation in the pine stem. II. Responses of differentiating tracheids

The effect of auxin supplied to the main stem of 5-year-old <em>Pinus silvestris</em> trees during various periods after decapitation upon differentiation of the secondary xylem tracheids was investigated. The results revealed the complexity of auxin involvement in the regulatory system of tracheid differentiation of secondary xylem. It is manifested both as the inductive effect to which the cells respond in the meristematic phase and in the continuous control during the consecutive stages of radial growth and maturation. A lack of auxin during the meristematic phase resulted in smaller cell diameters and reduced the daily rate of cell wall deposition even though these cells progressively grew and matured in the presence of auxin. The intensity of these two processes increased and the cells deposited thicker walls when auxin was supplied during all stages of tracheid differentiation even though the period of maturation decreased. Under these conditions tracheids of compression wood type differentiated. Continuous availability of auxin causes earlier termination of tracheid maturation while lack of auxin results in a delay of autolysis of protoplasts. In this case auxin probably functions in a system specifying information concerning the position of the cells in respect to the cambial layer.

Xuanweioxylon scalariforme gen. et sp. nov.: Novel Permian coniferophyte stems with scalariform bordered pitting on secondary xylem tracheids

Fossil lignophyte stems show remarkable diversity including non-sympodial and sympodial forms and a variety of pith and tracheid structures. Most lack scalariform bordered pits on radial walls of secondary xylem tracheids, while these occur in the putative progymnosperm Protopitys and some (but not all) cycads, bennettitaleans, and angiosperms. We describe a new genus of Permian gymnosperms that comprises decorticated stems and branches with a parenchymatous pith that is septate in stems but entire in branches and has scleroid cells that are organised singly or in bands. Vasculature is non-sympodial, comprising endarch primary xylem strands that become mesarch leaf traces that are occluded through ontogeny. Secondary xylem is pycnoxylic with scalariform bordered pits on radial tracheid walls. The eustelic and non-sympodial organisation, the septate pith and the presence of scalariform bordered pits on secondary tracheids represent a unique combination of features not previously seen in lignophyte genera, justifying the erection of Xuanweioxylon scalariforme gen. et sp. nov. X. scalariforme shows similarities to several extinct and extant lignophytes but is placed within the coniferophytes — an informal grouping of conifer-like plants of uncertain affinity that includes probable sister groups to cordaitaleans and conifers. Scalariform bordered pitting in secondary tracheids is unique to Xuanweioxylon within coniferophytes, demonstrating previously unrecognised diversity within the group. Results suggest that coniferophytes include intermediates between cordaitaleans and conifers with other lignophytes including Protopitys, pteridosperms, cycads and Bennettitales.

A polyxylic Cycad trunk from the Middle Jurassic of western Liaoning, China, and its evolutionary implications

A new fossil cycad stem Sinocycadoxylon liianum gen. et sp. nov. is described based on a well-preserved petrified specimen collected from the Middle Jurassic of Beipiao, Liaoning Province, China. The stem is composed of a small pith, a polyxylic vascular cylinder and cortex. The pith is parenchymatous with medullary vascular bundles, transfusion cells and mucilage canals. The polyxylic vascular cylinder consists of sixteen centrifugal rings and one centripetal vascular ring, all with distinct growth rings in secondary xylem. The presence of polyxyly with growth rings distinguishes this new genus from all fossil and extant Cycadales. The primary xylem is mesarch. Secondary xylem tracheids have araucarioid radial pitting. The rays are uni- or multiseriate and the latter are usually accompanied by mucilage canals and transfusion cells. This polyxylic stem is believed to be related to the Encephalartoideae of Zamiaceae. The new find provides new insights into cycad evolution which is consistent with former phylogenetic analyses indicating that Cycadales originated form Medullosales.

Reconstruction of the Pennsylvanian-age walchian conifer Emporia cryptica sp. nov . (Emporiaceae: Voltziales)

This paper provides a whole plant concept for a new species, Emporia cryptica sp. nov. Hernandez-Castillo, Stockey, Rothwell & Mapes (Emporiaceae: Voltziales), the fourth conifer to be reconstructed from the rich fossil biota at the Late Pennsylvanian, Hamilton Quarry, Kansas. E. cryptica has an orthotropic stem, lateral plagiotropic branches with simple leaves, simple pollen cones, and compound ovulate cones. Branches have an endarch eustele with dense wood surrounding a parenchymatous pith with sclerotic nests/plates, and secondary xylem tracheids with multiseriate hexagonal bordered pits. Leaves on both penultimate and ultimate branches are simple and amphistomatic with two adaxial stomatal bands, monocyclic and dicyclic stomata, and two narrow abaxial rows of stomata with numerous trichome bases. Pollen cones are simple with helically arranged microsporophylls and adaxial pollen sacs. Prepollen is monolete, eusaccate, and monosaccate ( Potonieisporites Bharadwaj). Ovulate cones are compound with bilaterally symmetrical dwarf shoots in the axils of helically arranged forked bracts. Axillary dwarf shoots bear numerous sterile scales interspersed with two megasporophylls, each bearing a single inverted terminal ovule. This plant displays morphological and cuticular characters similar to several morphogenera of Paleozoic walchian conifers but is most comparable to the fossil plant species Emporia lockardii and Hanskerpia hamiltonensis. E. cryptica is the only walchian conifer where ovules and seeds with megagametophytes, immature embryos and mature embryos have been documented, demonstrating that the most ancient conifers possessed seed dormancy and polycotyledonary embryos. This reproductive biology is similar to that of many Mesozoic and extant conifers with saccate pollen except for the presence of prepollen grains that are common among Paleozoic walchian conifers.

A WOODY LYCOPSID STEM FROM THE NEW ALBANY SHALE (LOWER MISSISSIPPIAN) OF KENTUCKY

A lycopsid axis from the New Albany Shale (Sanderson Formation) of Kentucky is described. The stem, which branches dichotomously, is 45 mm in diameter and is characterized by a relatively narrow parenchymatized protostele, a 3.0 mm‐thick cylinder of secondary xylem, a tripartite cortex, and a periderm that is more than 5.0 mm thick. The secondary xylem is composed of uniseriate and biseriate vascular rays and narrow tracheids with scalariform wall thickenings on both radial and tangential walls. The periderm is characterized by elongate, thick‐walled cells, some of which broaden tangentially in the outer part of the tissue forming zones that appear wedge‐shaped in cross section. Surface features of the axis, including leaf bases, are not preserved. The stem is tentatively regarded as a member of the Lepidodendrales in accordance with the numerous anatomical characters that it shares with more recent representatives of the order. Because the external morphology is not known, however, the possibility exists that the axis corresponds to a protolepidodendralean taxon currently known only from compression and/or impression remains or some other nonlepidodendralean plant that produced secondary xylem. The extremely narrow profile of the secondary xylem tracheids (relative to other arborescent lycopsids) is interpreted as evidence that the plants grew in a habitat that was substantially drier than the Upper Carboniferous coal swamps.

STRUCTURALLY PRESERVED FOSSIL PLANTS FROM ANTARCTICA. I. ANTARCTICYCAS, GEN. NOV., A TRIASSIC CYCAD STEM FROM THE BEARDMORE GLACIER AREA

Silicified stems with typical cycadalean anatomy are described from specimens collected from the Fremouw Formation (Triassic) in the Transantarctic Mountains of Antarctica. Axes are slender with a large parenchymatous pith and cortex separated by a narrow ring of vascular tissue. Mucilage canals are present in both pith and cortex. Vascular tissue consists of endarch primary xylem, a narrow band of secondary xylem tracheids, cambial zone, and region of secondary phloem. Vascular bundles contain uni‐ to triseriate rays with larger rays up to 2 mm wide separating the individual bundles. Pitting on primary xylem elements ranges from helical to scalariform; secondary xylem tracheids exhibit alternate circular bordered pits. Traces, often accompanied by a mucilage canal, extend out through the large rays into the cortex where some assume a girdling configuration. A zone of periderm is present at the periphery of the stem. Large and small roots are attached to the stem and are conspicuous in the surrounding matrix. The anatomy of the Antarctic cycad is compared with that of other fossil and extant cycadalean stems.

Initiation of adventitious root primordia in very young Pinusbanksiana seedling cuttings

The present work describes the anatomy of adventitious root initiation in 20-day-old Pinusbanksiana Lamb, seedling cuttings propagated under intermittent mist. Shortly after cuttings were made, basal necrosis occurred in all tissues (epidermis, periderm, cortex, primary and secondary phloem, and vascular cambium) that surrounded the central xylem cylinder. Thereafter, a relatively small "callus complex" composed of parenchyma cells, a few secondary xylem tracheids, and incompletely differentiated callus vascular cambium and periderm developed at the base of cuttings. One or sometimes two root primordia initiated in the transition zone between the lowermost cortical cells of the hypocotyl and the uppermost callus parenchyma cells. Primordia invariably arose just outside one of the four axial resin canals in the hypocotyl. Results suggested that adventitious root primordia may be initiated in P. banksiana cuttings only in association with differentiated or differentiating resin canals.