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SEASONALITY in CAMBIAL ACTIVITY of FOUR LIANAS from a MEXICAN LOWLAND TROPICAL RAINFOREST Calixto Leon-Gomez1 & Arcadio Monr

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SEASONALITY in CAMBIAL ACTIVITY of FOUR LIANAS from a MEXICAN LOWLAND TROPICAL RAINFOREST Calixto Leon-Gomez1 & Arcadio Monr IAWA Journal, Vol. 26 (1), 2005: 111-120 SEASONALITY IN CAMBIAL ACTIVITY OF FOUR LIANAS FROM A MEXICAN LOWLAND TROPICAL RAINFOREST Calixto Leon-Gomez1 & Arcadio Monroy-Ata2 SUMMARY The rhythm of vascular cambial activity was studied in four species of lianas growing in a lowland tropical rainforest of Mexico. Cambial activity was determined by counting the number of layers of cells in the cambial zone. A greater number of layers was assumed to indicate greater cambial activity. In all four species (Machaerium cobanense, M.floribundum, Gouania lupuloides and Trichostigma octandrum) the cambium is active throughout the year. In three of the species (all but T. octandrum) cambial activity was significantly higher in the April-October rainy period than in the November-March dry period. Cambial activity in Trichostigma octandrum was not significantly associated with the wet or dry season. Key words: Cambial activity, successive cambia, tropicallianas, tropi­ cal rainforest. INTRODUCTION Most studies of cambial activity have been conducted in trees, with lianas only rarely being considered. The first studies of cambial activity in lianas, inc1uding seasonal changes in cambial activity, were on temperate species such as Vitis vinifera, V. riparia, Parthenocissus inserta and Celastrus scandens, and the tropical Bougainvillea glabra (Esau 1948; Davis & Evert 1970; Pulawska 1973). Other workers have investigated seasonal changes in lianas by focusing on phenology (Croat 1975; Putz & Windsor 1987; Hegarty 1988; Opler et al. 1991). These studies show the dearth of information on lianas, and the need to understand the correlation between environmental factors and stern structure. In tropical and subtropical regions, correlation of cambial activity with seasons is not well understood for most species and there is often no c1ear separation of pronounced pe­ riods of activity and rest. In some tropical trees the cambium can remain active during the entire year, with peak periods correlated with rapid shoot extension. But in general, litde is known about the patterns of activity in tropical trees, inc1uding both the initiation and cessation of growth, due to the fact that very few studies have been conducted in tropical regions. As could be expected, little is also known about how lianas grow. 1) Departamento de Botanica, Instituto de Biologfa, UNAM, A. P. 70-233, Mexico 04510, D. E, Mexico. 2) Unidad de Investigaci6n en Ecologfa Vegetal, FES Zaragoza, UNAM, A. P. 9-020, Mexico 15000, D.E, Mexico. Associate Editor: Peter Gasson Downloaded from Brill.com10/02/2021 04:16:13PM via free access 112 IAWA Journal, Vol. 26 (I), 2005 In many trees, growth rings suggest periodicity in increase in diameter of the stern, and though this phenomenon is little understood in tropical trees, growth rings are commonly observed. Growth rings are rare in lianas, making their age determination difficult. Therefore, this paper attempts to document the pattern of cambial activity in four species of liana from the tropical rainforest of the Los Tuxtlas region, Veracruz, Mexico. This area is mo ist year-round, with a peak during the single yearly rainy season. The temperature remains relatively constant throughout the year. The main objective of this paper is to test the hypothesis that lianas grow continuously throughout the year but with peaks in growth during the rainy season. MATERIALS AND METHODS Study area - Los Tuxtlas tropical biology field station is on the slopes of the Gulf of Mexico, in the southeast part of the State of Veracruz, with an elevation of 150 to 530 m, between 95° 04' and 95° 09' West longitude and between 18° 34' and 18° 36' North latitude (Lot-Helgueras 1976) (Fig. 1).The climate is warm-humid [Af(m)w"(i') following the classification of Köeppen modified by Garcfa 1981], with an annual pre- 95"10' 95"00' 18"40' 18"40'_ San Marlin Tuxtla ~ Volcano ~ 18"30' 18"30'- Santiago Tuxtla 18"20'_ o 5 10 I I km Felipe Villegas 1L/1995 95"00' Fig. 1. Map of the study area. The Tropical Biology Station is situated in southeast Mexico, at 25 km from the Catemaco-Montepio road. Downloaded from Brill.com10/02/2021 04:16:13PM via free access Le6n-G6mez & Monroy-Ata - Cambial activity in lianes 113 1400 2 1200 1000 E .§.. 800 ~ 600 ';;" "" 400 200 0 F M A M A o N 0 Month. 35 3 a 30 , - ~ -- <! , ~ 25 -.. E . ,! ... .. .~ ... - .... - 8- ' -.~ - E ,.-:.. - .~ .., .-: .... b ~ 20 ~. '- '''.':'''.,;",.' 15 J M A M A 0 Monlh Fig. 2 & 3. Precipitation and temperature graphs from the "Los Tuxtlas" station. The continuous line corresponds with the year 1998; the dashed line with 1999 and the dotted line with 2000. a: maximum temperatures; b: minimum temperatures. cipitation of 4700 mm (Fig. 2). The maximum temperature is 32.3 oe and the minimum temperature is 16.4 oe, with a mean of 24.3 oe (Fig. 3). Plant material - The Iianas chosen for this study are characteristic of the primary forest of the region and reach basal stern diameters of up to 20 cm. The species inves­ tigated are Machaerium cobanense Donn.Sm. and M.floribundum Benth (Fabaceae), Gouania lupuloides (L.) Urban (Rhamnaceae) and Trichostigma octandrum (L.) H. Walt. (Phytolaccaceae) (Fig. 4). Monlhs: F D Flowcring: FrUClifi al ion: •••• Vegelillivc growlh: •••• Fig. 4. Species phenogram (Machaerium cobanense, M.floribundum, Trichostigma octandrum, Gouania lupuloides). The flowering norrnally occurs at the end of the dry period and the fruc­ ti/ication appears at the beginning of the rainy period. Machaerium cobanense is out of phase. Downloaded from Brill.com10/02/2021 04:16:13PM via free access 114 IAWA Journal, Vol. 26 (1), 2005 Field work - The study took place from January 1999 through December 2000. For each species, two individuals with a stern diameter of 1-5 cm were collected monthly. A section of stern 2 cm long was taken from each individual and preserved in FAA (Sass 1958). When the sampies were less than 2.5 cm in diameter, transections were made of the entire stern. When the sampies exceeded 2.5 cm in diameter, they were sub-sampled in 6 to 8 transections containing vascular cambium and adjacent tissues. Microtechnique - The xylem of these species is tough, especially in sterns of more than 1 cm in diameter, so the sampies were softened in a heated solution of 10% ethy­ lene-diamine for 30-60 min (cf. Kukachka 1977; Carlquist 1982), and embedded in polyethylene glycol. Sections of 15-20 !Am were cut with a sliding microtome. Sections were stained with safranin-aniline blue (Johansen 1940). Determination ofcambial activity - Cambial activity was determined by counting the number of layers or radial files of cells in the cambial zone. A greater number of lay­ ers of cells was assumed to indicate greater cambial activity (Paliwal & Prasad 1970: p. 337; Siddiqi 1991: p.179;Larson 1994: p. 597; Priya& Bhat 1999: p.183). Thetenns "cambium" and "cambial zone" are used in this study to designate the complete meri­ stematic zone including the fusifonn cambial initials and part ofthe phloem and xylem, of which the cells are enlarging but still capable of dividing, as described by Wilson et al. (1966). For each of the two individuals sampled monthly, from 4 to 8, but nonnally 6 counts of cambiallayers were made. For the transections of whole sterns, counts were made in four equidistant points, and for the subsectioned sterns, one count was made in each sub-sample. At-test was used to compare within-species differences between the rainy and the dry seasons for each species. RESULTS Phenology and structure of the cambium The species studied are evergreen but have smaller leaf crowns during the dry season. Machaerium ftoribundum, Gouania lupuloides and Trichostigma octandrum mainly flower in the January-April dry season. However, Machaerium cobanense flowers in the rainy months of May to August (Fig. 2 & 4). All species nonnally fmit at the beginning of the rainy season, from March to July. Development of secondary xylem in the four species includes two types of cambia. Gouania lupuloides and Trichostigma octandrum have a 'nonnal' cambium in which the secondary xylem is produced as a continuous cylinder, whereas Machaeriumftori­ bundum and M. cobanense have successive cambia, in which the xylem alternates with concentric rings of phloem, corresponding to the Securidaca type of Carlquist (1988). In transection, the fusifonn and ray initials distinguish them from the cambial derivatives because of the abundance of periclinal divisions and thinner walls. The organization as viewed in tangential sections of the cambium varies for each species: no storying was observed in T. octandrum, while M. cobanense, M.ftoribundum and G.lupuloides have ray and parenchyma cells storied (Fig. 5). Downloaded from Brill.com10/02/2021 04:16:13PM via free access Le6n-G6mez & Monroy-Ata - Cambial activity in lianes 115 Fig. 5. Tangential stern seetions that show the longitudinal organization of xylem elements. - A : Machaerium cobanense. - B: M.jioribundum. - C: Gouania lupuloides. - D: Trichostigma octandrum. - M. cobanense, M. jioribundum and G. lupuloides have storied rays and paren­ chyma strands, while in T. octandrum there was no storying. - ap = storied axial parenchyma; r =storied ray. - Scale bar =200 ILm. Annual variation in cambial activity In all four species, the vascular cambium is active all year round (Fig. 6 & 8), but the number of cambial celllayers is at its maximum during the rainy period. In Mach­ aerium cobanense, development is at its maximum of 4 to 6 celllayers during Febru­ ary to October with a mean number of layers of 3 to 4 the rest of the year (Fig. 8A). Machaeriumfloribundum presents a maximum of 4 to 7layers of cambium in July and August (Fig. 8B); Gouania lupuloides is at its maximum from April to October, with the formation of 6 to 8layers of cells, while from November to March only 4 to 5 layers are observed (Fig.
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