Anatomical Investigation of Root, Stem and Branch Wood in 10-Year-Old Inga Laurina in the Context of Anatomical Adaptation to Hydraulic and Mechanical Stresses
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Anales de Biología 40: 31-39, 2018 ARTICLE DOI: http://dx.doi.org/10.6018/analesbio.40.04 Anatomical investigation of root, stem and branch wood in 10-year-old Inga laurina in the context of anatomical adaptation to hydraulic and mechanical stresses Eduardo L. Longui1, Ana T. D. Galão1, Kishore S. Rajput2 & Antônio C. G. de Melo1 1 Instituto Florestal, CEP 02377-000, São Paulo, SP, Brazil. 2 Department of Botany, Faculty of Science, The Maharaja Sayajirao University of Baroda, Vadodara 390002, India. Resumen Correspondence Investigación anatómica de la madera de la raíz, el tallo y la rama E.L. Longui en ejemplares de 10 años de edad de Inga laurina en el contexto E-mail: [email protected] de la adaptación anatómica al estrés hidráulico y mecánico Received: 5 September 2017 La madera varía entre las especies, desde la médula hasta la cor- Accepted: 5 February 2018 teza, desde las raíces hasta la corona dentro de la misma planta, Published on-line: 7 March 2018 para garantizar la seguridad y la eficiencia hidráulica. La heteroge- neidad de la madera de Inga laurina (Sw.) Willd. es aquí investiga- da, siendo una especie que prospera bien en todos los bosques brasileños importantes. La madera de tallo de I. laurina poseía va- sos estrechos; fibras de paredes gruesas; rayos más anchos, altos y en mayor frecuencia, características que proporcionan ajustes hi- dráulicos y mecánicos. Las fibras de paredes gruesas y menos pa- rénquima, en comparación con la raíz, brindan la fuerza suficiente para apoyar la corona grande. Una mayor proporción de células de parénquima en las raíces da como resultado una gran capacidad de almacenamiento de almidón, que es importante para la recupe- ración de los brotes después del daño por fuego, común en el Ce- rrado. Palabras clave: Cerrado, Conductividad hidráulica, Densidad, Estructura arbórea, Madera tropical o subtropical. Abstract Wood varies between species, from pith to bark, roots to crown within the same plant to ensure hydraulic safety and efficiency. Inga laurina (Sw.) Willd. is investigated herewith for the structural heterogeneity of the wood, since this species thrives well in all ma- jor forests of Brazil. I. laurina stem wood possessed narrow ves- sels; thick walled fibres; wider, taller and higher frequency of rays, features that provide hydraulic and mechanical adjustments. Thick walled fibres and less parenchyma as compared to root provide enough strength to stem to support the large crown. Higher propor- tion of parenchyma cells in roots results in a high storage capacity for starch, which is important for the shoot recovery after fire dam- age, common in the Cerrado. Key words: Cerrado, Hydraulic conductivity, Density, Tree structure, Tropical or subtropical wood. 32 E.L. Longui et al. Anales de Biología 40, 2018 Introduction variation in the hydraulic conductivity and wood density of I. laurina wood at different positions Inga laurina (Sw.) Willd. (Fabaceae) is a native viz. root, main stem and branch. We hypothesize species of Brazil (Pennington 1997, Garcia 1998). that I. laurina, a species with a large crown is able It is distributed from north to south of Amazon, in to adjust its stem wood to perform necessary func- Caatinga, Cerrado and Atlantic Rainforest. It also tions that are associated with hydraulic conductiv- grows naturally in Rupestrian Field, Semi-decidu- ity. However, available literature indicates that ous Forest, Ombrophylous Forest and Restinga trees are able to regulate one parameter without (Garcia & Fernandes 2014). Typically, this species affecting the other that is associated with former is found on river margins or on dry fields, as well one (Awad et al. 2012, Badel et al. 2015). as interfluve regions of the Cerrado (Pennigton 1997) and in gaps in the Amazonian Forest (Souza Material and methods et al. 2011). In all these forests, amount of annual precipitation varies from region to region, yet the Study area present species thrives well in all these forest types. The study was carried out in the Cerrado area of the Assis State Forest (22º39”42’S, 50º24”44’ W), Structurally, the secondary xylem is heteroge- Assis, São Paulo (Max et al. 2007). According to neous (i.e. non-uniform); particularly variation in Bognola et al. (2003), the soil in this region has xylem structure from pith to bark, root to apical low organic matter with good permeability and meristem, and from stem to branch. Therefore, good water holding capacity. The climate is Cwa this heterogeneity in the secondary xylem struc- by the Köppen climate classification: warm with ture results in spatial variation in hydraulic and mechanical performance of the xylem (Gartner dry winters, mean annual temperature of 20 °C, 1995). For instance, concerning hydraulic effi- mean maximum of 25 °C and average minimum is ciency and mechanical strength trade-off, high of 18 °C. The average annual rainfall is 1441.5 frequency of large vessels has positive influence mm (CEPAGRI 2017). The region has a dry sea- on hydraulic efficiency and negative influence on son between April and September (see Fig. 1). mechanical strength (Baas et al. 2004). In con- Sampling trast, variation in abiotic and biotic environments often compels trees to respond through growth Transplantation of I. laurina planting was esta- rate like height, stem diameter, canopy cover, and blished in 2002 at a distance of 3 m x 2 m. After root depth (Hinckley et al. 2011) to compete for 10 years, 11 trees were felled and circular discs of above and below ground resources. the root at 0.3 m depth from ground level and Recent studies are focused on the structural above ground samples were obtained from two variation in the axial conducting elements of different positions i.e. at 0.5 m (stem) and 1.5 m plants, which attempt to explain how xylem struc- i.e. from main branch (branch which forked more ture provides hydraulic safety and water conduct- centrally in relation to the stem) in height (Fig. 2 ing efficiency from roots to leaves for plants and Table 1). growing under higher salinity, water limitation or Wood anatomy, hydraulic conductivity and high levels of rainfall variability, and in drought- density tolerant conditions (Robert et al. 2009, von Arx et al. 2012, Abrantes et al. 2013, Anfodillo et al. Small wood pieces (outermost 2-3 yrs. growth 2013). increment) adjacent to bark were cut from the four Such findings open up a vast field for new different sides of the disc that were obtained from research into the axial variations of vessels and roots, stem and branches. Thin slices of wood tracheids a as water conducting cells and varia- material were obtained with the help of razor tions in fibers and parenchyma cells that together blade and macerated according to the modified forms the wood structure. Thus, it is challenging Franklin method (Berlyn & Miksche 1976) to to the wood researchers to focus their attention on obtain dimensional details of the individual wood plant species living under different conditions, elements. Thirty measurements were taken ran- whether in natural forests or artificial forest plant- domly to obtain mean and standard deviations. ings. Present study anatomically investigated the For sectioning, as mentioned above, outermost 2- Anales de Biología 40, 2018 Wood axial variation of Inga laurina 33 Tree nº 1 2 3 4 5 6 7 8 9 10 11 Tree height (m) 6.2 6.2 6.4 6.2 6.3 6.0 5.5 5.7 5.2 5.8 6.3 ø 1.5 m (cm) Main branch 8.5 7.1 9.9 9.7 9.0 7.2 5.0 6.4 4.7 7.8 7.5 ø 0.5 m (cm) Stem 11.8 11.6 12.6 11.7 12.5 10.5 7.9 8.8 6.3 9.1 12.4 ø Root (cm) 30 cm depth 3.6 5.3 4.6 3.8 9.8 5.7 4.9 8.3 5.8 4.0 4.7 Tabla 1. Altura del árbol y diámetro de la raíz, tallo y rama principal de Inga laurina con 10 años de edad. Table 1. Tree height and diameter of root, stem and main branch of 10-year-old Inga laurina. Figura 2. A: Árbol de Inga laurina y muestreo; B: Ilustración esquemática que muestra los puntos de extracción de las muestras de madera para su investigación. Figure 2. A: Inga laurina tree and sampling; B: Schematic illustration showing the points of removal of wood samples for investigation. The terminology and characterization of wood is followed as per the IAWA list (IAWA Committee Figura 1. A: Localización del Bosque Estatal de Assis (*) en el 1989). All anatomical measurements were Estado de São Paulo, Brasil; B: Precipitación media mensual, obtained using an Olympus CX 31 microscope déficit hídrico DEF(-1) (abril y agosto), excedente hídrico EXC y temperatura media en el Bosque Estatal de Assis desde 1961a equipped with a digital camera (Olympus Evolt E 1990 (CEPAGRI, 2017). 330) and image analyzer software (Image-Pro Figure 1. A: Location of Assis State Forest (*) in São Paulo 6.3). State, Brazil; B: Mean monthly precipitation, water deficit DEF(- 1) (April and August), water surplus EXC and mean temperature Vessel grouping index was calculated as at Assis State Forest from 1961-1990 (CEPAGRI, 2017). described by Carlquist (2001) and according to the equation of Scholz et al. (2013): Vg = N ves- sels/N groupings, where Vg = vessel grouping and 3 year’s wood increments were cut from the four groupings = solitary, multiples of 2, 3 and multi- opposite sides of all 11 wood disks each that were ples of 4 or more vessels.