Báo cáo khoa học: " Vegetative development, primary and secondary growth of the shoot system of young Terminalia superba tropical trees, in a natural environment. I. Spatial variation"

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Original article Vegetative development, primary and secondary growth of the shoot system of young Terminalia superba tropical trees, in a natural environment. I. Spatial variation in structure and size of axes E de Faÿ Université de Nancy I, Laboratoire de biologie des Ligneux, BP 239, 54506 Vand&oelig;uvre-lès-Nancy Cedex, France (Received 21 August 1991; accepted 17 April 1992) Summary — Spatial variations in trunk structure including wood histology and size of axes were ex- amined in 5 21-month-old Terminalia superba Engl and Diels trees, grown in a natural tropical envi- ronment. Long and shorter internode series alternated along the main axes, forming rather poorly delimited units of extension. In trunk wood, the paratracheal axial parenchyma of the confluent type formed irregular bands, the spacing of which varied gradually, increasing and decreasing alternately. These widely- and narrowly- spaced parenchyma bands differentiated various wood layers. The radi- al enlargement and wall thickness of fibres also varied in the same manner, as did, but to a lesser extent, the width of parenchymal bands, the radial enlargement of parenchyma cells and vessels, and the wall thickness of vessels. The number of various wood layers in trunk segments that were defined by variations in intemodal length was not related to the order of appearance of the trunk seg- ments in question. These results suggest that trunk wood exhibited alternating dense and not very dense wood layers, but they did not provide visible structural evidence of a well-pronounced growth periodicity in the main shoot of Terminalia superba. The existence of a structural discontinuity in the peripheral wood and the significance of the basipetal differentiation of the outermost adjacent wood layer have been discussed in relation to seasonal changes in climate and phenology. On the other hand, trunk thickness increased markedly from 5 cm above to 5 cm beneath the upper pseudo- whorls of branches. The thickness of axes (branches and trunks) was closely related to the number of sympodial units located in a centrifugal and an upper position respectively. Moreover, a pseudo- whorl of branches was generally inserted in the middle part of trunk segments defined by variations in internodal length. These data indicate trunk-branch correlations. It is suggested that some struc- tural variations in trunk wood could be related to branching. radial size / structural unit / Terminalia superba / tropical tree / wood Résumé — Développement végétatif, croissance primaire et secondaire du système cauli- naire de jeunes arbres tropicaux de l’espèce Terminalia superba, dans un environnement na- turel. I. Variation dans l’espace de la structure et de la dimension des axes. Les variations dans l’espace de la structure du tronc - y compris l’histologie du bois - et de la dimension des axes furent examinées chez 5 plants de Terminalia superba Engl et Diels âgés de 21 mois et poussant dans un environnement naturel tropical. Des séries d’entre-n&oelig;uds longs et plus courts alternaient le long des
principaux formant des unités de croissance mal délimitées. Dans le bois des troncs, le paren- axes chyme axial paratrachéal, de type confluent, formait des bandes irrégulières dont l’espacement variait graduellement, augmentant et diminuant en alternance. Ces bandes de parenchyme largement et fai- blement espacées différenciaient des couches de bois. La dilatation radiale et l’épaisseur des parois des fibres variaient aussi de la même manière que précédemment. Il en était de même, quoique à un moindre degré, de l’épaisseur des bandes de parenchyme, de la dilatation radiale des cellules paren- chymateuses et des vaisseaux, et de l’épaisseur des parois des vaisseaux. Le nombre de couches de bois dans les segments de tronc définis par des variations dans la longueur des entre-n&oelig;uds n’était pas lié à l’ordre d’apparition des segments en question. Tous ces résultats suggèrent que le bois des troncs présentait des couches alternativement denses et peu denses, mais ils ne donnent pas de preuve structurale d’une périodicité de croissance bien marquée dans la flèche des Terminalia superba. L’existence d’une discontinuité structurale dans le bois périphérique et la signification de la couche de bois adjacente la plus externe sont discutées en relation avec les changements de saison et de phénologie. Par ailleurs, l’épaisseur des troncs s’accroissait de façon marquée de 5 cm au- dessus à 5 cm en-dessous des étages supérieurs de branches. L’épaisseur des axes (branches et troncs) était étroitement liée au nombre d’unités sympodiales en position respectivement centrifuge et supérieure. De plus, un étage de branches était généralement inséré dans la partie médiane des seg- ments de tronc définis par des variations dans la longueur des entre-n&oelig;uds. Ces données attirent l’attention sur les corrélations entre branches et tronc. II est suggéré que des variations structurales dans le bois du tronc pourraient être liées à la ramification. dimension radiale / unité structurale /Terminalia superba / arbre tropical / bois INTRODUCTION growth among the trees of the inter- cambial However, detailed studies tropical zone. on cambial activity, such as histophysiological Some attention has been paid to growth investigations are still rare, fragmentary and phenomena in tropical trees. Morphological generally concern mature trees growing in- and morphogenetic studies on shoot growth termittently, the wood of which displays an- have demonstrated the basic knowledge of nual growth rings (Lowe, 1968; Paliwal and continuous and flushing - or rhythmic - Prasad, 1970; Lawton and Lawton, 1971; growth and architectural models (Bond, Rao, 1972; Amobi, 1973; Paliwal et al, 1942; 1945; Scarrone, 1965; Hallé and 1976; Ghouse and Shamima Hashmi, 1979; Martin, 1968; Borchert, 1969; Hallé and Zamski, 1979; Rao and Dave, 1981; Rog- Oldeman, 1970; Greathouse et al, 1971; ers, 1981; Dave and Rao, 1982; Desh- Vogel, 1975a,b; Hallé et al, 1978; Ng, 1979; pande and Rajendrababu, 1985; Venugopal Parisot, 1988; El-Morsy and Millet, 1989). and Krishnamurthy, 1987). As regards cor- Concerning secondary growth, there is relations between growing organs, only re- some information on wood production and lationships between leaf and shoot growth on the existence, anatomy and periodicity have been examined in a few tropical trees of growth rings in the wood of numerous (Greathouse et al, 1971; Prévost, 1972; species (Hummel, 1946; Mariaux, 1967; Borchert, 1973, 1978) and the effect of Lowe, 1968; Mariaux, 1969, 1970; Gill, leaves on internode elongation has been 1971; Amobi, 1974; Détienne et Mariaux, demonstrated in Terminalia superba (Mail- 1975, 1976, 1977; Fahn et al, 1981; Mari- lard et al, 1987b). aux, 1981; Zamski, 1981; Ash, 1983a, b; de Faÿ, 1985; Boninsegna et al, 1989; Dé- Young plants of the African species Ter- minalia superba Engl and Diels grown in a tienne, 1989; Jacoby, 1989; Worbes, 1989). These data suggest various patterns of natural environment studied within were
the context of research MATERIALS AND METHODS on growth rhythms and relationships between primary and secondary growths of the shoot system. T The trees studied came from the experimental superba is a forest species, the architec- plantation of the Centre Technique Forestier ture of which has been characterized by Tropical on the Ivory Coast, located in the forest Hallé and Oldeman (1970) as Aubréville’s of Anguédédou, about 30 km north-west of Abid- jan. A clonal plot was put at the author’s dispo- model. Its trunk is a monopode, the growth sal to study 5 trees. The trees were descended of which is assumed to be rhythmic and from stem cuttings that had been made from 3- gives branches a pseudowhorled arrange- month-old shoots and had been planted 3 ment, commonly called a tier. Branches months later. They were aged 21 months when are sympodial and horizontal by apposition studied. A good size, well-separated tiers of to spiral phyllotaxy units. Each sympodial branches and an absence of any trunk reitera- tion were the criteria for the choice of plants. unit is characteristic of the genus Terminal- The clonal origin was unknown at the time of se- ia, the Terminalia branching being specifi- lection, which should have ensured a certain cally described in T catappa (Hallé and variability among the chosen trees, as if seed- Oldeman, 1970; Tomlinson and Gill, 1973; lings had been studied. The 5 trees (TA-TE) Fisher, 1978). The vegetative development were of the same geographic provenance, 2 of T superba, at an early stage, has recent- were half-sibs (TA and TC) and 2 were of the same clone (TD and TE). The ground had been ly been studied under controlled conditions prepared, before planting, by manual cutting of (Maillard, 1987; Maillard et al, 1987a,b; the forest plot; the plant spacing was initially 2.5 1989). These authors determined the most x 2.5 m; manual weeding had been regular. favourable conditions for growth and con- Therefore, the T superba trees were not growing sidered growth periodicity. It appears that in competition for light with other vegetation. the trunk displays an alternation of long They had been subjected to full sunlight. and short internode series, obvious at As the species T superba is disseminated 27 °C and under 14- or 16-h daylengths. throughout the evergreen rain forest in the south of the Ivory Coast and it invades the secondary This paper reports a study on the spatial bush (Aubréville, 1959), the variability and local aspects of growth in a natural tropical envi- growing conditions of the trees studied were ap- parently similar to those of ’fraké’ saplings - fra- ronment. The purpose was not only to com- ké is the local common name of this species - in pare the of young vegetative development a large chablis of the adjacent forest. The site is trees under natural and controlled environ- located in the intertropical zone at about 5° lati- mental conditions, especially the formation tude north, that is to say in the so-called ’humid of morphologically discrete growth incre- in many tropical regions, the season- tropics’. As ments (units of extension, Hallé et al, 1978), al variations in temperature and solar radiation are not great, but the amount of rainfall is distrib- but also to increase the number of criteria uted irregularly. There are 2 dry seasons (De- which would allow the growth of Terminalia cember-March and mid July-September) alter- superba to be characterized. Trunk wood nating with 2 rainy seasons (April-mid July and structure and size of axes were thus investi- mid September-November). Thus for more than gated. On the one hand, relationships be- one and a half years, the young trees had been tween radial size of axes and number of subjected to a monthly temperature of 25-28 °C branch modular components, ie sympodial (Atlas de la Côte d’Ivoire, 1978) and a photoperi- od of 12.2-13.2 h daylength, allowing for dawn units, were sought in addition to units of ex- and dusk (Longman and Jenik, 1963), which is tension along trunks. On the other hand, fairly close to the 27 °C and 14-h daylength histological investigations on trunk wood found to be the most favourable for the develop- were aimed at finding structural variations in ment of young T superba trees under controlled relation to units of extension, ie rhythmic conditions (Maillard et al, 1987a). However, they growth rings (de Faÿ, 1985). had lived through several dry seasons and the
last one was just before the examination. The at the served the trunks, except along top; lack of rainfall could thus have restricted plant only a sharp limit at the indeed, there was growth seasonally. base of the developing part, in 4 out of 5 For observations and measurements, trunks trees. However, on the lower part, there cut at the base, at the beginning of March, were several areas with some relatively were just after total new leafing of the plants, and short internodes, compared with the adja- were inspected immediately from base to apex. cent regions. Variations in internodal The relative position of leaf-scars, or leaves in length occurred progressively. It should be the upper part only, was studied to find regions noted that a slight variation might pass un- with short internodes or an alternation of long and shorter internode regions. The transition noticed and, moreover, leaf-scars tended zones from long to short internode regions were to become indistinct on the old parts of marked to delimit units of extension. When the trunks. Nevertheless, trunk segments con- limit was not obvious, the middle of the shortest sisting of long internode series separated internode was taken into account to separate by shorter internode series could be distin- successive trunk segments, and these were guished (fig 1). measured. In addition, the number of tiers per unit, the number of branches per tier, the num- These poorly delimited segments gener- ber of sympodial units per branch, the mean ba- ally bore one tier of branches located in sal diameter of branches and the circumference the central part. However, 2 sets of of trunks at 5 cm above and below each tier, or observed in 2 different seg- branches in the middle of a unit with no branch, were re- were corded. and, in the basal segments, ments some- times branch found For (fig 1). histological studies of the wood, 2 kinds no was of sample were collected in the middle part of Concerning axis sizes, the circumfer- each unit of extension, beneath the tier of of trunks increased from apex to ence branches: discs and cores. Firstly, the 2-3-cm base but irregularly on each side of the de- long discs were put into an air-conditioned room veloping tiers of branches. In addition, the to dry; afterwards, they were treated with sodi- um hypochlorite to lighten them (Mariaux, 1969) basal diameter of branches was correlated and finely polished; they were observed with a positively with the number of component stereomicroscope before and after treatment sympodial units (fig 2). A close relationship with phloroglucinol-HCl, characterizing lignins was also found between trunk circumfer- (Sass, 1958). Secondly, the outermost trunk tis- ence and the total number of sympodial including wood, were sampled with a sues, units in the branches inserted above the bone puncture surgical instrument and the cores were fixed in Craf 1 (Sass, 1958); they level studied (fig 3). were then cut transversely with a freezing mi- crotome, and the 40-μm thick sections were stained with phloroglucinol-HCl and examined Structural variations in trunk wood microscopically. The wood of the trunks showed slight con- RESULTS centric colour variations in polished trans- verse sections. A stereomicroscope high- lighted numerous light irregular concentric Spatial variations bands on a comparatively dark back- in external structure and size of axes ground (figs 4A, B). These bands were paratracheal axial parenchyma of the con- Typical units of extension, ie trunk seg- fluent type. In general, band spacing var- ments, well-delimited by areas of more ied gradually, increasing and decreasing alternately. The dark background of the tightly packed leaf-scars, could not be ob-
transverse sections of wood con- polished sisted of wood fibres. Positive phlorogluci- nol-HCl tests showed wood fibres ar- ranged in alternating concentric layers (fig 5A). These variations resulted mainly from changes in the number of cells per unit area (fig 5B). These 2 types of concentric and repeat- ed variations - in the spacing of parenchy- ma bands and in the intensity of staining - were not always closely correlated, espe- cially in the lower trunk segments. More- over, the thickness of wood layers varied strongly within a trunk section, particularly in the lower segments: a single wood layer was sometimes thick and sometimes thin; the different wood layers that were defined by the same feature had very different thicknesses, in the internal wood, whatever their position; the thickness of adjacent wood layers varied independently: for in- stance the wood layers with relatively nar- row band spacing were sometimes thick and sometimes thin in comparison with the innermost ones with wider band spacing. The great structural variability in trunk wood made the wood layer count difficult. Several counts of the wood layers that were defined by variations in band spacing were made in each trunk sections (on the 2 polished areas), because a small local variation of band spacing could at first pass unnoticed. The results (table I) showed clearly that the number of wood layers increased basipetally but irregularly from one trunk segment to the next (older one) below. The most recent wood layers (2-5 ac- cording to the trunk level) were distin- guished from each other more easily: those with a wide spacing of parenchyma bands were thick, whereas those with nar- row band spacing were thin; moreover, parenchyma bands were wider in the for- mer than in the latter (figs 4A, B). The staining of the wood background following
the walled fibres, the former belonging to the phloroglucinol-HCl test was less in- tense in the wood layers with wide band outermost wood layer that was defined by spacing than in those with narrow band narrowly-spaced parenchyma bands and spacing. Variations in staining were gradu- the latter to the differentiating wood which al from one wood layer to another, except exhibited widely-spaced parenchyma bands between the 2 most recent wood layers (fig 6B). This boundary was not always uni- (fig 6A). form at the cellular level, mainly because of the occurrence of parenchyma bands. In Indeed, the limit was sharp only be- this area, parenchyma cells changed from tween the 2 most recent wood layers. His- a radially flattened type to a radially dilated tologically, this was the transition between type - more precisely from a rather narrow strongly radially flattened, thick-walled fi- area to a wider area in cross section (fig bres and younger radially dilated, thinner-
and vessels from small, thick-walled by alternating series of long and shorter in- 6B) - pores to large, relatively thin-walled pores controlled environment in ternodes, as a (fig 6A). The microscopical examination chamber (Maillard et al, 1987a), which did not indicate noticeable changes in wall generally formed poorly-delimited units of staining. Histological differences between extension. The vaguely segmented struc- wood layers with narrow and wide spac- ture of the trunk was confirmed at the ings of parenchyma bands tended to be wood level. The spacing of parenchyma bands, the radial enlargment of wood fi- less visible in internal wood (figs 4A, B, 5A, B). The developing wood layer was found bres and parenchyma cells, the wall thick- ness of wood fibres and to a lesser extent in the 5 trunks studied, but its thickness the width of parenchyma bands, the pore decreased basipetally and it could be par- size and wall thickness of vessels changed tially formed or totally missing in the base in such a way that wood layers were sug- of the trunk (figs 4A, B). gested to be alternately more and less dense. The changes were generally gradu- DISCUSSION al and apparently bore no relation to the development of the units of extension. The trunk of young T superba trees grown Thus, there were no rhythmic growth rings in the juvenile wood of T superba, unlike in a natural environment was characterized
that of Hevea brasiliensis (de Faÿ, 1985; rings of mature tropical (Mari- trees some 1986). In short, structural units of the He- 1967; Boninsegna et al, 1989; Dé- aux, vea type, characteristic of the rhythmic tienne, 1989; Vetter and Botosso, 1989; growth-habit were found neither externally Worbes, 1989). In fact, the abrupt (spatial) along the trunk nor in trunk wood. variations of trunk thickness were related to the upper tiers of branches and there Nevertheless, the structural variations was a close relationship between trunk in wood could indicate that the cambial thickness and size of branches (in number growth of trunks was alternately rapid and of modules) that were inserted above the slow, especially since structural variations of that nature are visible within the growth level studied. Moreover, trunk segments
that were defined by variations in inter- al, 1971; Vogel, 1975a, b; de Faÿ, unpub- nodal length usually bore 1 tier of branch- lished Quercus robur results) (Payan, or es. These data indicate trunk-branch cor- 1982; Champagnat et al, 1986). These re- relations and imply that some structural sults on spatial variations in structure and variations in trunk wood could be related to size of axes indicate trunk-branch correla- branching. Indeed, the leaves of develop- tions rather than typical units of extension along the trunk and rhythmic growth rings in ing axes are assumed to produce the stim- uli (phytohormones) that induce the differ- trunk wood. A subsequent study of temporal entiation of wood (Aloni, 1989). aspects of growth in young T superba trees in the same natural environment (de Faÿ, Climatic variations could also have had 1992) has already given results in accor- repercussions on wood formation, espe- dance with the present data. It is thus sug- cially the lack of rainfall, as in the wood of gested that the ’pagoda’ architecture of Ter- Carya glabra where the spacing of tangen- minalia superba resulted more from the tial parenchyma bands is related to the periodicity of branching than from a flushing rate of rainfall (Hill, 1982). At any rate, the growth of trunk, at least in the early stage. outer structural discontinuity in the trunk wood of the trees studied could be related to a seasonal change, at least in phenolo- ACKNOWLEDGMENTS gy. Indeed, initiation of the basipetally de- veloping wood layer was associated in time with the bud break (this had begun 2 The author is grateful to the late Director of the weeks before sampling and had followed Centre Technique Forestier Tropical on the Ivo- ry Coast, Dr K Diabate and his colleagues for the totally leafless period occurring at the providing plant material and Professor JM Favre end of the long dry season). Reduction or (University of Nancy I) for his critical reading of absence of the developing wood layer at the manuscript. the base of trunks should indicate a delay in the resumption of cambial activity, be- cause of the basipetal decrease in leaf hor- REFERENCES mone stimulation (Aloni, 1989). The peri- pheral structural discontinuity in trunk Aloni R (1989) Control of xylogenesis within the wood should be the sign of a rest during whole tree. Ann Sci For, 46 suppl 267s-272s the totally leafless period when the trees Amobi CC (1973) Periodicity of wood formation were almost 2 years old. It should be point- in some trees of lowland rainforest in Nigeria. ed out that some younger T superba plants Ann Bot 37, 211-218 growing in the same climate showed low or Amobi CC (1974) Periodicity of wood formation no radial increments, or trunk shrinkage in in twigs of some tropical trees in Nigeria. Ann February, ie at the end of the long dry sea- Bot 38, 931-936 son (de Faÿ, 1992). Wood differentiation in Ash J (1983a) Growth rings in Agathis robusta the trunks of young T superba plants could and Araucaria cunninghamii from tropical thus be altered by drought other or causes Australia. Austr J Bot 31, 269-275 of internal water deficit. (1983b) Tree rings in tropical Callitris ma- Ash J In conclusion, the trunk of young T su- cleayana F Muell. Aust J Bot 31, 277-281 perba tropical trees grown in a natural en- Aubréville A (1959) La Flore Forestière de la vironment did not have the typical features Côte d’Ivoire. Centre Tech For Trop, Nogent of flushing species, such as Hevea brasi- Marne, France, vol 3 sur liensis (Hallé and Martin, 1968; de Faÿ, Bond TET (1942) Studies in the vegetative 1985), Theobroma cacao (Greathouse et growth and anatomy of the tea plant (Camel-
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