Segments Of Cambium Research Articles

Ontogeny of multiple variants in the stems of Phaseolus lunatus L. (Fabaceae)

Development of the cambial variant is characteristic to most of the climbing members of the Fabaceae and is considered as an alternative path to increase their stem diameter for efficient supply of minerals and photosynthate within limited stem diameter. These alternative forms of secondary growths are ascribed as an adaptive character to increase stem flexibility and most often they are found to be taxon-specific. In the present study, histological investigation on the main stems of Phaseolus lunatus L. (Fabaceae) showed multiple combinations of variant secondary growth like the formation of vascular cylinders external to the secondary phloem and proliferation of parenchyma (axial and ray) followed by the development of interxylary phloem, interxylary cambia (functionally regular and inverse) and ray cambia. The secondary xylem produced by regular cambium was diffuse-porous with indistinct growth rings and composed of an abundance of thin-walled unlignified parenchyma. With the increase in age, phloem rays underwent dilatation, subsequently became meristematic and formed oval to circular vascular cylinders/units with irregular orientation. Concomitantly, dedifferentiation xylem parenchyma not only formed small islands of interxylary phloem but also led to the origin of interxylary cambial segments that differentiated bidirectionally and deposited xylem and phloem having inverse, diagonal, tangential or radial orientations. Similar cell division activity was also observed in marginal xylem ray cells (ray cambium) that produced both the xylem and phloem on either side. All these cambial variants are described in detail and their possible significance in the climbing habit is discussed.

Development of successive cambia and structure of secondary xylem in the stems and roots of Distimake tuberosus (Convolvulaceae)

In the present study, development of trilobed stem is investigated in Distimake tuberosus (Convolvulaceae) by histological methods. Increase in stem thickness and formation of trilobed stem was achieved by forming crescent shaped successive cambial segments from the pericyclic parenchyma at three equidistant sites. In contrast, the alternate segments of the vascular cambium were functionally less active, dividing leisurely (referred henceforth as functionally less active cambium) and did not form the successive cambium. The newly formed cambial segments divide bi-directionally and produced its derivatives, while functionally less active segment lagged behind. The secondary xylem formed from the functionally less active cambial segments have unique pattern of vessels distribution i.e., wide vessels rare or completely absent. Parenchyma (both, axial and ray) proliferation, formation of ray cambium, functionally inverse cambia, tyloses in fibriform vessels and the occurrence of bordered pits in xylem ray cells are observed in all samples. Root diameter increased by developing concentric rings of successive cambia and maintained a circular outline. Formation of successive cambia on three sides and alteration in stem outline may be associated with protection of conducting elements against stem torsion and internal injury. Formation of conducting elements from the ray cambium and internal cambium suggest rapid and safe conduction of photosynthate, water and mineral elements.

Development of successive cambia and formation of secondary xylem in Suaeda nudiflora and S. fruticosa (Amaranthaceae s.l.)

Amaranthaceae s.l. and its allied families like Amaranthaceae s.s., Chenopodiaceae s.s. and Phytolaccaceae s.s. are always debated in the literature about their pattern of secondary growth; when their stem diameter increases by forming successive cambia. In the present study, development of successive cambia and differentiation of its derivatives was investigated in Suaeda fruticosa and S. nudiflora (Amaranthaceae s.l.). After primary growth, the first complete ring of cambium was formed by the successive cambium due to failure in the development of interfascicular cambium between adjacent vascular bundles. The first ring of successive cambium initiated from the parenchymatous cells (pericyclic derivatives) external to the protophloem. This newly formed cambium was functionally bidirectional and produced secondary xylem centripetally and phloem centrifugally. Subsequent renewal of cambium was observed as small segments instead of complete ring. They interconnect with existing cambium and form anastomosing network. Due to the renewal of small sectors, the secondary phloem formed by earlier cambial segments became enclosed within the secondary xylem and conjunctive tissue. Formation of vessels and sieve elements were found confined only to few fusiform cells while rest of the cambial sector exclusively produced conjunctive tissue on either side. The secondary xylem formed in the beginning of successive rings of xylem remained rayless while thick stems showed presence of heterocellular rays with several cells in height and width. Accumulation of starch along with the presence of nuclei in the xylem fibres even after deposition of the secondary wall is consistent feature in both the species.

Development of successive cambia and structure of the secondary xylem in some members of the family Amaranthaceae

Young stems of Aerva javanica (Burm.f.) Juss. ex Schult., A. lanata (L.) Juss. ex Schult, A. monsonia Mart., A. sanguinolenta (L.) Blume, Alternanthera bettzickiana (Regel) G. Nicholson, A. philoxeroides (Mart.) Griseb., Gomphrena celosioides Mart., G. globosa L. and Telanthera ficoidea (L.) Moq., showed the renewal of small sectors of cambium by replacing with new segments. Therefore, the secondary phloem formed by earlier cambial segments form isolated islands of phloem enclosed within conjunctive tissues became embedded in the secondary xylem. As the stem grows older, complete ring of cambium is renewed; sometimes an anastomosing network of successive cambia may be seen due to the renewal of larger segments of the cambium. Renewal of the cambium takes place by repeated periclinal division in the parenchyma cells positioned outside to the phloem formed by the previous cambium. Functionally the cambium is bidirectional and exclusively composed of fusiform cambial cells. Differentiation of conducting elements of the secondary xylem and phloem remains restricted to the certain cambial cells while rest of the segments exclusively produce conjunctive cells. Accumulation of starch along with the presence of nuclei in the xylem fibers even after deposition of the secondary wall is consistent in all the species and it seems to be associated with the absence of rays in the secondary xylem and phloem of nine species from four genera. The significance of successive cambia, rayless xylem and nucleated xylem fibers were correlated with plant habit.

Development of successive cambia and formation of flat stems in Rhynchosia pyramidalis (Lam.) Urb. (Fabaceae)

Stem flattening in Rhynchosia pyramidalis (Fabaceae) is achieved by the development of crescent-shaped successive cambia on two opposite sides of the stem (referred hereafter as distal side). Other lateral sides of the stem (adjacent to supporting host and its opposite side, referred as proximal sides) usually possess single cambium. In the young stems, parenchymatous cells located outside to protophloem of distal side dedifferentiate and develop small segments of cambium. Concomitant to bidirectional differentiation of the secondary xylem and phloem, these newly developed cambial segments also extend in tangential directions. Differential activity of newly developed crescent-shaped cambial segments deposits more secondary xylem at median position as compared to their terminal ends of the stem on distal side; consequently, it pushes the cambial segment outside, thus resulting in crescent-shaped arcs of the cambia only on two opposite sides. After the production of 1–2 mm of secondary xylem, they cease to divide and new segments of cambial arc develop on the same side in a similar fashion. Such repeated behaviour of successive cambia development consequently leads to the formation of tangentially flat stems. The secondary xylem is diffusely porous with indistinct growth rings and is composed of vessels (wide and narrow), fibres, axial ray parenchyma cells, while phloem consisted of sieve elements, companion cells, axial and ray parenchyma. Rays in both xylem and phloem are uni- to multiseriate and heterocellular. The structure of secondary xylem and development of successive cambia is correlated with climbing habit.

Internal cambium and intraxylary phloem development in Ipomoea turbinata Lag. (Convolvulaceae)

Strands of phloem that are present at the periphery of pith are known as intraxylary phloem. Its presence remains restricted to a small portion of eudicots and considered as a characteristic feature for certain families. In the present study, Ipomoea turbinata Lag. showed development of intraxylary protophloem on adaxial tips of protoxylem from the procambial derivatives. Subsequently, additional intraxylary sieve elements were added from the adjacent parenchyma cells that were morphologically different from the pith cells. In thick stems, thin walled cells located between protoxylem and intraxylary protophloem acquired meristematic characters and formed several small segments of internal cambium. Initially, these cambial segments produced only phloem derivatives. Soon after, these segments became bidirectional and began to produce secondary xylem centrifugally and secondary phloem centripetally. Interestingly, in some of the samples development of secondary phloem and xylem was observed in the same direction. The secondary xylem formed by the internal cambium was composed of wide and fibriform vessels, fibres and axial parenchyma. Phloem possessed sieve tube elements, companion cells and parenchyma cells while rays in both xylem and phloem were mostly uniseriate but multiseriate rays were also observed.

Stem anatomy and development of inter- and intraxylary phloem in Leptadenia pyrotechnica (Forssk.) Decne. (Asclepiadaceae)

Modification of external morphology and internal structure of plants is a key feature of their successful survival in extreme habitats. They adapt to arid habitats not only by modifying their leaves, but also show several modifications in their conducting system. Therefore, the present study is aimed to investigate the pattern of secondary growth in Leptadenia pyrotechnica (Forssk.) Decne., (Asclepiadaceae), one such species growing in Kachchh district, an arid region of Gujarat State. A single ring of vascular cambium, responsible for radial growth, divided bidirectionally and formed the secondary xylem centripetally and the phloem centrifugally. After a short period of secondary xylem differentiation, small arcs of cambium began to form secondary phloem centripetally instead of secondary xylem. After a short duration of such secondary phloem formation, these segments of cambium resumed their normal function to produce secondary xylem internally. Thus, the phloem strands became embedded within the secondary xylem and formed interxylary phloem islands. Such a recurrent behavior of the vascular cambium resulted in the formation of several patches of interxylary phloem islands. In thick stems the earlier formed non-conducting interxylary phloem showed heavy accumulation of callose on the sieve plates followed by their crushing in response to the addition of new sieve elements. Development of intraxylary phloem is also observed from the cells situated on the pith margin. As secondary growth progresses further, small arcs of internal cambium get initiated between the protoxylem and intraxylary phloem. In the secondary xylem, some of the vessels are exceptionally thick-walled, which may be associated with dry habitats in order to protect the vessel from collapsing during the dryer part of the year. The inter- and intraxylary phloem may also be an adaptive feature to prevent the sieve elements to become non-conducting during summer when the temperature is much higher.

Development of successive cambia and wood structure in stem of Rivea hypocriteriformis (Convolvulaceae)

Abstract This study examined the formation of successive rings of cambia in Rivea hypocriteriformis Choisy (Convolvulaceae). The mature stem is composed of four to five rings of xylem alternating with phloem. Successive cambia originate as smaller and larger segments; union and anastomosing of small cambial segments often leads to the formation of discontinuous rings. In the initial stage of growth, several vascular bundles interconnect to form the first ring of vascular cambium. The cambium remains functional for one complete season and becomes dormant during summer; a new ring of cambium is completed prior to the subsequent monsoon season and sprouting of new leaves. Successive cambia are initiated from the pericyclic parenchyma situated three to four cell layers outside of the protophloem. Functionally, all the successive cambia are bidirectional and produce secondary xylem centripetally and phloem centrifugally. The secondary xylem is diffuse-porous, with indistinct growth rings and consisting of wide fibriform vessels, fibre tracheids, and axial and ray parenchyma cells. The xylem rays are uni- to multiseriate and heterocellular. The multiseriate rays contain lignified marginal ray cells and thin-walled, unlignified central cells. The central ray cells also show accumulations of starch and druses. Discrete strands of intraxylary phloem occur at the periphery of the pith, and additional intraxylary phloem develops from adjacent cells as secondary growth progresses. Earlier-formed phloem shows heavy accumulation of callose, followed by its compaction. The development of successive cambia is correlated with extension growth and with the phenology of the plant.

Development of Successive Cambia and Structure of Secondary Xylem of Ipomoea Obscura (Convolvulaceae)

Abstract Stems of Ipomoea obscura Ker Gawl., increase in thickness by forming multiple rings of cambia. Stems 5-6 mm thick produce parenchymatous derivatives which divide repeatedly to form small arcs of cambium. Several such small arcs initiate simultaneously and form a ring of small cambial arcs. After the formation of a few xylem and phloem elements, all these arcs are interconnected by transdifferentiation of parenchyma cells present between the cambial arcs and constitute a complete cambial cylinder. This newly formed cambium is functionally bidirectional: earlier- formed arcs produce xylem centripetally and phloem centrifugally, while later-formed segments exclusively produce thin-walled parenchyma cells on either side. Young stems are circular in cross section but as stem thickness increases they become oval to elliptic or lobed and dumbbell-shaped. Xylem rays are mostly uni- or biseriate and thin-walled, but multiseriate rays characteristic for a climbing habit are observed occasionally. In thick stems, the marginal ray parenchyma in most of the samples becomes meristematic and develops ray cambia which exclusively produce sieve elements. Similarly, parenchyma cells produced from later-formed cambial segments give rise to several irregularly oriented vascular bundles. The secondary xylem is diffuse porous, with indistinct growth rings and is composed of fibriform and wider vessels, fibres, and axial and ray parenchyma cells, while phloem consists of sieve elements, companion cells, and axial and ray parenchyma cells.

Development of Intra- and Interxylary Secondary Phloem in Coccinia indica (Cucurbitaceae)

Stem anatomy and the development of intraxylary phloem were investigated in six to eight years old Coccinia indica L. (Cucurbitaceae). Secondary growth in the stems was achieved by the normal cambial activity. In the innermost part of the thicker stems, xylem parenchyma and pith cells dedifferentiated into meristematic cells at several points. In some of the wider rays, ray cells dedifferentiate and produce secondary xylem and phloem with different orientations and sometimes a complete bicollateral vascular bundle. The inner cambial segments of the bicollateral vascular bundle (of primary growth) maintained radial arrangement even in the mature stems but in most places the cambia were either inactive or showed very few cell divisions. Concomitant with the obliteration and collapse of inner phloem (of bicollateral vascular bundles), parenchyma cells encircling the phloem became meristematic forming a circular sheath of internal cambia. These internal cambia produce only intraxylary secondary phloem centripetally and do not produce any secondary xylem. In the stem, secondary xylem consisted mainly of axial parenchyma, small strands of thick-walled xylem derivatives, i.e. vessel elements and fibres embedded in parenchymatous ground mass, wide and tall rays along with exceptionally wide vessels characteristic of lianas. In thick stems, the axial parenchyma de-differentiated into meristem, which later re-differentiated into interxylary phloem. Fibre dimorphism and pseudo-vestured pits in the vessels are also reported.

Formation of successive cambia and stem anatomy ofSesuvium sesuvioides(Aizoaceae)

Mature stems of Sesuvium sesuvioides (Fenzl) Verdc. were found to be composed of successive rings of xylem alternating with phloem. Repeated periclinal divisions in the parenchyma outside the primary phloem gave rise to conjunctive tissue and the lateral meristem that differentiate into the vascular cambium on its inner side. After the formation of the vascular cambium, the lateral meristem external to it became indistinct as long as the cambium was functional. As the cambium ceased to divide, the lateral meristem again became apparent prior to the initiation of the next cambial ring. The cambium was exclusively composed of fusiform cambial cells with no rays. In the young saplings, the number of cambial cylinders in the axis varied from the apex to the base, indicating formation of several rings within the year. In each successive ring of the lateral meristem, small segments differentiated into the vascular cambium and gave rise to vessels, axial parenchyma, fibres and fibriform vessels towards the inside, and secondary phloem on the outer side. In the old stems, non-functional phloem of the innermost rings was replaced by a new set of sieve tube elements formed by periclinal divisions in the cambial segments associated with the non-functional phloem. In some places the cambial segments completely differentiate into derivatives leaving no cambial cells between the xylem and phloem. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 158, 548‐555.

Radial secondary growth and formation of successive cambia and their products inIpomoea hederifoliaL. (Convolvulaceae)

Ipomoea hederifolia stems increase in thickness using a combination of different types of cambial variant, such as the discontinuous concentric rings of cambia, the development of included phloem, the reverse orientation of discontinuous cambial segments, the internal phloem, the formation of secondary xylem and phloem from the internal cambium, and differentiation of cork in the pith. After primary growth, the first ring of cambium arises between the external primary phloem and primary xylem, producing secondary phloem centrifugally and secondary xylem centripetally. The stem becomes lobed, flat, undulating, or irregular in shape as a result of the formation of both discontinuous and continuous concentric rings of cambia. As the formation of secondary xylem is greater in one region than in another, this results in the formation of a grooved stem. Successive cambia formed after the first ring are of two distinct functional types: (1) functionally normal successive cambia that divide to form secondary xylem centripetally and secondary phloem centrifugally, like other dicotyledons that show successive rings, and (2) abnormal cambia with reverse orientation. The former type of successive rings originates from the parenchyma cells located outside the phloem produced by previous cambium. The latter type of cambium develops from the conjunctive tissue located at the base of the secondary xylem formed by functionally normal cambia. This cambium is functionally inverted, producing secondary xylem centrifugally and secondary phloem centripetally. In later secondary growth, xylem parenchyma situated deep inside the secondary xylem undergoes de-differentiation, and re-differentiates into included phloem islands in secondary xylem. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 158, 30–40.

Stem anatomy of Dolichos lablab Linn (Fabaceae): Origin of cambium and reverse orientation of vascular bundles

Secondary growth in the stem of Dolichos lablab is achieved by the formation of eccentric successive rings of vascular bundles. The stem is composed of parenchymatous ground tissue and xylem and phloem confined to portions of small cambial segments. However, development of new cambial segments can be observed from the obliterating ray parenchyma, the outermost phloem parenchyma and the secondary cortical parenchyma. Initially cambium develops as small segments, which latter become joined to form a complete cylinder of vascular cambium. Each cambial ring is functionally divided into two distinct regions. The one segment of cambium produces thick-walled lignified xylem derivatives in centripetal direction and phloem elements centrifugally. The other segment produces only thin-walled parenchyma on both xylem and phloem side. In mature stems, some of the axial parenchyma embedded deep inside the xylem acquires meristematic activity and leads to the formation of thick-walled xylem derivatives centrifugally and phloem elements centripetally. The secondary xylem comprises vessel elements, tracheids, fibres and axial parenchyma. Rays are uni-multiseriate in the region of cambium that produces xylem and phloem derivatives, while in some of the regions of cambium large multiseriate, compound, aggregate and polycentric rays can be noticed.

Initiation and structure of the secondary vascular system in Phytolacca dioica (Phytolaccaceae)

Phytolacca dioica (L.) is characterized by anomalous secondary thickening by means of supernumerary cambia. After a period of primary growth and the formation of an initial (normal) vascular cambium, supernumerary cambia are initiated outside of the primary vascular cylinder. The initiation of the first supernumerary cambium takes place through approximately the number of nodes equal to the denominator of the phyllotactic fraction characterizing a given axis. At each node a segment of supernumerary cambium is initiated opposite the leaf traces supplying the leaf inserted at that node. The segments of differentiated cambium are preceded by regions of obliquely and anticlinally dividing cells. In the single juvenile axis studied supernumerary cambial segments also appear above the node to the cathodic side of the entering leaf traces, and opposite the medullary bundle immediately anodic to these traces. Vascular connections among the primary and supernumerary vascular cylinders occur between leaf insertions on the same orthostichy. The levels at which these connections occur vary among stems. The switch from ordinary to anomalous secondary growth may be caused by a change in tissue response to stimuli produced by leaves.

ONTOGENY AND CORRELATIVE RELATIONSHIPS OF THE PRIMARY THICKENING MERISTEM IN FOUR‐O'CLOCK PLANTS (NYCTAGINACEAE) MAINTAINED UNDER LONG AND SHORT PHOTOPERIODS

The developmental anatomy of Mirabilis jalapa was investigated during the first 90 days of growth. The primary thickening meristem (PTM) initially differentiates in the pericycle at the top of the cotyledonary node 18 days after germination, then basipetally in the pericycle through the hypocotyl. The PTM differentiates acropetally into the stem and in the pericycle of the primaiy root, commencing 22 days after germination. Endodermis is easily identifiable in hypocotyls as well as in primary roots because of Casparian thickenings in its cells. It has not been definitely identified in stems. There are three rings of primary vascular bundles in the stem. The PTM differentiates as segments of cambium in a layer of cells (probably in the pericycle) on an arc between vascular bundles of the outer bundle ring. Later, arcs of PTM differentiate externally to the phloem of each bundle. Each arc forms a connection between original segments of PTM lying on either side of each vascular bundle. Thus, the PTM becomes a continuous cylinder. The PTM differentiates in the pericycle outside vascular tissue in the hypocotyl and root. Differentiation of the PTM and the mode of secondary thickening is similar in plants exposed to short (8‐hr) and to long (18‐hr) photoperiods, but some differences were observed. The PTM differentiates closer to the stem apex in all plants over 18 clays of age growing vegetatively under long photoperiods. That is, the diffuse lateral meristem, in whose cells the PTM differentiates in young intemodes, is shorter in nearly all investigated plants growing in long photoperiods. The hypocotyl and base of the primary root of 40‐day‐old plants in short photoperiods were more enlarged than those of the same age plants in long photoperiods; but, at the end of 64 days, the hypocotyl and primaiy root base were larger in plants growing under short photoperiods. Thirty‐four days after seed germination, flower initiation occurs in plants exposed to short photoperiods. One hundred fifty days after seed germination, flowers differentiate on plants exposed to long photoperiods.