Coppice Polar Architecture

Tree Physiology 23, 11531170 2003 Heron PublishingVictoria, Canada
A method for describing the canopy architecture of coppice poplar with allometric relationships
ERIC CASELLA1,2 and HERV SINOQUET3
1 2 3
Downloaded from http://treephys.oxfordjournals.org at University of Novi SadFaculty of Law on June 14, 2010
Forest Research, Mensuration, Alice Holt Lodge, Farnham GU10 4LH, U.K. Author to whom correspondence should be addressed (eric.casella@forestry.gsi.gov.uk) UMR Physiologie Intgre de lArbre Fruitier et Forestier, INRA-Universit Blaise Pascal, Site de Croul, 234 avenue du Brzet, 63039 Clermont-Ferrand cedex 02, France
Received March 23, 2003; accepted May 24, 2003; published online November 3, 2003
Summary A multi-scale biometric methodology for describing the architecture of fast-growing short-rotation woody crops is used to describe 2-year-old poplar clones during the second rotation. To allow for expressions of genetic variability observed within this species (i.e., growth potential, leaf morphology, coppice and canopy structure), the method has been applied to two clones: Ghoy (Gho) (Populus deltoides Bartr. ex Marsh. Populus nigra L.) and Trichobel (Tri) (Populus trichocarpa Torr. & A. Gray Populus trichocarpa). The method operates at the stool level and describes the plant as a collection of components (shoots and branches) described as a collection of metameric elements, themselves defined as a collection of elementary units (internode, petiole, leaf blade). Branching and connection between the plant units (i.e., plant topology) and their spatial location, orientation, size and shape (i.e., plant geometry) describe the plant architecture. The methodology has been used to describe the plant architecture of 15 selected stools per clone over a 5-month period. On individual stools, shoots have been selected from three classes (small, medium and large) spanning the diameter distribution range. Using a multi-scale approach, empirical allometric relationships were used to parameterize elementary units of the plant, topological relationships and geometry (e.g., distribution of shoot diameters on stool, shoot attributes from shoot diameter). The empirical functions form the basis of the 3-D Coppice Poplar Canopy Architecture model (3-D CPCA), which recreates the architecture and canopy structure of fast-growing coppice crops at the plot scale. Model outputs are assessed through visual and quantitative comparisons between actual photographs of the coppice canopy and simulated images. Overall, results indicate a good predictive ability of the 3-D CPCA model. Keywords: biometry, canopy structure, fish-eye lens, Populus deltoides Populus nigra, Populus trichocarpa, seasonal dynamics, short-rotation woody crop.
Introduction Populus spp. is one of the most promising fast-growing tree species for producing fuel and fiber on short-rotation coppice systems where crops are harvested every 3 years over a 15-year lifespan (Grassi et al. 1990, Macpherson 1995, Ceulemans et al. 1996). Large genotypic differences have been demonstrated among natural and hybrid clones in productivity (Heilman and Stettler 1985, Pontailler et al. 1999, Laureysens et al. 2000, Tubby and Armstrong 2002), leaf area index (LAI) (DeBell et al. 1996, Heilman et al. 1996, Casella and Ceulemans 2002) and other growth characteristics (e.g., plantsoilnutrient interactions, site climate, pests and diseases) (Liu and Dickmann 1992a, 1992b, Stettler et al. 1996, Ceulemans and Deraedt 1999, Dickmann et al. 2001). Poplar taxa differ considerably in canopy architecture and seasonal dynamics through shoot demography (Laureysens et al. 2000), leaf phenology (Ceulemans et al. 1988, Deraedt and Ceulemans 1998), leaf and branch morphology and spatial distribution (Ceulemans et al. 1990, Dunlap et al. 1992, Casella and Ceulemans 2002), and growth characteristics (Ridge et al. 1986, Ceulemans et al. 1988, Pieters et al. 1999). In particular, productive hybrid poplar clones differ from less productive natural ones in leaf size (maximum/average: 600/90 versus 120/20 cm2 for Populus trichocarpa Torr. & A. Gray Populus deltoides Bartr. ex Marsh. hybrid clone Hoogvorst and P. nigra L. clone Wolterson, respectively), leaf inclination angles (0 40 versus 0 80 on average for clones Hoogvorst and Wolterson, respectively), and the distribution of these foliar characteristics within the canopy (Ceulemans and Isebrands 1996, Eckenwalder 2001, Casella and Ceulemans 2002). Because plant productivity is directly related to the ability to intercept photosynthetically active radiation (e i) (see Table 1 for a list of symbols and their definitions) and convert it to biomass through photosynthesis (e c) (Cannell et al. 1988, Cannell 1989), leaf area density (LAD) and leaf attributes (i.e., locations, orientations, sizes and shapes) (Takenaka 1994) are key canopy parameters needed to describe the radiation regime within a canopy for simulating the mass and energy ex-
1154 Table 1. Symbols and abbreviations. Symbol A A a ab a10 a Cl db d10 dt DlI DnL d ec ei I Definition
CASELLA AND SINOQUET
l L LAD LAI M N n w P j y S Se q U
Axis. A1, an axis of order 1, is a shoot, and A2, an axis of order 2, is a branch Total leaf area per U2 (cm2) Individual L area (cm2) Basal stool section area (cm2) Shoot section area at 10 cm above the point at which the shoot joins the stool (cm2) Inclination angle between A1 and A2 or A2 and P () The stool or the shoot diameter class number Diameter at the bottom of a stool or shoot (cm) Diameter at 10 cm above the point at which a shoot joins a stool (cm) Diameter at the shoot tip (cm) Internode elongation growth potential Leaf initiation rate Branch or petiole divergence angle () Canopy light conversion efficiency Canopy light interception efficiency Internode. An internode is morphologically represented as a cylinder, bounded at its ends by nodes Length (cm) Leaf blade Leaf area density (m2 m3) Leaf area index Metameric unit Node. A node consists of a possibly vegetative bud, leaf or branch Number of Random deviation Petiole. A petiole is the slender stalk that supports the leaf blade Elevation angle of the normal to the vertical for I, Se and L () Twist angle of the normal to the vertical for L () Stool 20-cm long segment on A1U1 Azimuth angle of the normal () Growth unit. U1 is the part of an axis that has lengthened during the previous growing season, U2 is the growing part of an axis during the current year Width (cm)
change between a coppice crop and the atmosphere (Milne et al. 1992). The models that have been developed to simulate mass and energy exchange between plants and the atmosphere differ mainly in their treatment of canopy architecture (Ross 1981, Chen 1992, Chen et al. 1994a, 1994b, Law et al. 2001, Sinoquet et al. 2001). Because of difficulties in estimating the spatial distribution of leaf area within a canopy, the representation of the vegetation has generally been simplified. In the turbid medium approach (Ross 1981), canopies have been
abstracted as a leaf gas and divided into horizontal layers with uniform spatial distributions of the foliage (Law et al. 2001, Casella and Ceulemans 2002). But these one-dimensional models are unsuitable for horizontal or vertically heterogeneous or discontinuous canopies with narrowly or widely spaced plants and low leaf area in their earlier growth stages (such as forest landscape or short-rotation woody crop (SRWC) poplar systems) (Law et al. 2001, Casella and Ceulemans 2002). For these situations, canopy architecture can be described (i) as a collection of individual crowns modeled as 3-D geometric shapes (Law et al. 2001), (ii) by modeling the 3-D architecture of a population of plants using stochastic (De Reffye et al. 1988, 1991, Whitehead et al. 1990), fractal (Chen et al. 1994a) or L-system (Prusinkiewicz 1986, Prusinkiewicz et al. 1994) methods, or (iii) by describing accurately the geometry of each plant in situ, using the 3-D digitizing method (Sinoquet and Rivet 1997). The first approach has been successful in heterogeneous open-canopy forest landscape systems (Law et al. 2001). Nevertheless, this effort to consider the spatial heterogeneity of a canopy based on crown envelopes remains fundamentally one-dimensional at the crown or plot scales because the LAD within each foliage envelope is assumed to be uniformly distributed. The approaches based on stochastic, fractal or L-system theories improved the 3-D canopy architecture resolution by integrating both topological (Hall et al. 1978) and geometric (Ross 1981) notions of plant architecture. These methods can approximate many species shapes for studying lightvegetation interactions, but model outputs have never been precisely compared with field measurements (e.g., canopy openness). The digitizing method provides a precise description of the 3-D organization of every plant entity in space (i.e., from internode to leaf). Unfortunately, this method cannot be applied to poplar plants designed as SRWC systems because of their complex structure. To address this challenge, an alternate approach was to conceptualize the underlying plant architecture (after Godin et al. 1999) and use it as a guide for model construction and data acquisition. The objectives of this study were to (i) propose a multi-scale biometric methodology based on topological and geometrical information to deal with the 3-D architecture of fast-growing short-rotation poplar crops, (ii) apply the methodology to two hybrid clones of different genetic origin: Ghoy (Gho, Populus deltoides Populus nigra, characterized by a potential of productivity of about 4 Mg DM ha 1 year 1) and Trichobel (Tri, Populus trichocarpa Populus trichocarpa, characterized by a high potential of productivity of about 11 Mg DM ha 1 year 1), (iii) develop an empirical 3-D Coppice Poplar Canopy Architecture model (3-D CPCA), and (iv) evaluate the model quality from fish-eye photographs. Materials and methods Plant material and plantation layout Poplar plants were measured during the second growing season (2001) in a 0.5-ha experimental field in Farnham, U.K.
TREE PHYSIOLOGY VOLUME 23, 2003
CANOPY ARCHITECTURE OF COPPICE POPLAR Table 2. The two Populus clones used in this study. Name Code number Sex Parentage Parental code number S.9-2 Ghoy V.235 V.24 Provenance Latitude
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Longitude
Ghoy Trichobel
S.682-68 S.724-101
F M
P. deltoides P. nigra P. trichocarpa P. trichocarpa
Iowa, USA /Ontario, Canada Wallonnie, Belgium Washington, USA Oregon, USA
4249 N 51 N 49 N 4530 N
4 E 12230 W 12240 W
(5111 N, 051 E; 115 m a.s.l.). The plantation is situated on a stagnogley soil surrounded by forest, and consists of 16 different poplar (Populus spp.) clones belonging to different parentage and hybrid groups (Armstrong 1997). The clones Gho and Tri were used in this study (Table 2). A randomized block design with three replications for each clone was used, following the protocol prescribed by the U.K. Forestry Commission (Armstrong 1997). Hardwood cuttings (25 cm) were planted in April 1996 in double rows (oriented northsouth) with alternating inter-row distances of 0.75 m and 1.5 m (essential for mechanical harvests), and a spacing within rows of 0.9 m, to give a planting density of about 10,000 stools ha 1. Plots were 9 11.5 m and contained 10 rows of 10 stools each. In January 1997, stools were cut back to a height of 5 cm to create a multi-shoot coppice. For each clone, measurements were limited to 36 stools in the center of one plot. Descriptions of plant topology and geometry Two-year-old poplar plants were decomposed into components (stool, S; axis, A; growth unit, U; and metamer, M) and elementary units (internode, I; petiole, P; and leaf blade, L). On S, axes of order 1 (A1) were shoots connected to S, and axes of order 2 (A2) were branches connected to A1 (Figure 1). Growth unit 1 (U1) was the part of A1 that had lengthened during the previous growing season (i.e., 2000), and growth unit 2 (U2) was the growing part (or shoot leader) of A1 during the assessed year (i.e., 2001) (Figure 1). Because few lateral sylleptic A2 had lengthened on A1U1 during the previous growing season (i.e., A2U1), only proleptic A2 were described on A1U1 during the 2001 growing season (i.e., A2U2) (Figure 1). Each U was described as a succession of M, themselves defined as I bounded at their ends by nodes (N). Each N consisted of a vegetative bud, an A2U2 connected on A1U1, or an axillary bud + P + L connected on U2 (Figure 1). With this plant topology definition, 3-D plant architecture can be computer-generated if each elementary unit is given a shape, size, orientation and location in space. Basic geometric models (e.g., triangle, cylinder, frustum of a cone) are used to represent the shape (i.e., external surface) and size of plant units (e.g., the height of a triangle, l, and the width of its base, w). Detailed examples are available for tree architecture (Sinoquet and Rivet 1997, Godin et al. 1999), leaf geometry and canopy structure (Chen 1992, Sinoquet et al. 1998, Rakocevic et al. 2000, Dauzat et al. 2001, Sonohat et al. 2002).
In our study, S and P were represented as cylinders, U were divided into a sequence of conic frustums (i.e., I), and L were regarded as planar objects. A leaf blade prototype was created for each clone. Prototypes were represented as polygons with a set of four contiguous triangles for Clone Gho (Figure 1) and eight for Clone Tri, to fit the leaf blade shape and the allometric relationships between the leaf blade area (aL), the leaf midrib length (lL) and the leaf blade width (wL) as: aL = C l L wL (1)
where C is a regression coefficient. Finally, the representation of a basic geometric model in the scene refers to its position (i.e., orientation and location) with respect to a coordinate system (Figure 1). For L, its position in space is defined by the direction of the axis of symmetry of the polygon, or the leaf midrib, and the coordinates of the point of attachment of L to P (see Figure 1 for more details). Model description The model operates at the stool level and describes the plant architecture as a collection of A1. Each A1 is in turn described as a succession of components and elementary units defined by their size, shape, location, orientation and connection in space, which can change over the course of the growing season. From an input data file describing the locations (x and y) and basal diameters (d b) of stools in a vegetative scene, the modeling process starts by generating individual plants (i.e., stool + alive A1U1) as they were before bud burst in the spring of 2001. Each plant is then used as an initial fixed geometrical architecture (in terms of d b S; nA1U1, the number of A1U1 connected to a stool; lA1U1, the A1U1 length; nM, the number of metamers of a A1U1; and qA1U1, the A1U1 azimuth) for the scene reconstruction over the growing season (by describing the size, shape and position of each component carried by each M on each U). At the U and the leaf scales, the mortality was assumed to be negligible during the entire growing season. Initial plant architecture reconstruction The number of A1U1 connected to S is calculated from d b S as: nA1U1 = f ( d b S) (2)
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d10 A1U1i = 2
ab S i f p nA1U1
(3)
where abS is the basal stool section area and i = 1, , nA1U1 is the A1U1 number. From d10 A1U1, lA1U1 and nM are calculated as: lA1U1 = f ( d10 A1U1) nM = f (lA1U1) (4)
(5)
After numbering the M from the bottom to the top of A1U1, the elementary I units are described by their l, basal (d b) and top (d t ) diameter, according to the relative M rank (i/nM) and i -1 i the relative M position (1 l I j lA1U1 or 1 l I j lA1U1) on A1U1, as: i l I i = lA1U1 f nM
i -1 lI j d b I i = f d10 A1U1, 1 lA1U1 i lI d t I i = f d10 A1U1, 1 j lA1U1
(6)
(7)
(8)
Figure 1. Multi-scale codification of a poplar plant topology and description of Euler angles (q, azimuth; d, divergence; j, elevation; a, inclination; and y, twist) for assessing and computer-generating the three-dimensional plant architecture of 2-year-old poplar within highdensity coppice crops. Plant components and elementary units are stool (S), internode (I), petiole (P) and leaf blade (L). At a given scale, elementary units (e.g., I) of a plant component (e.g., axis of order 1 growth unit 1, A1U1) are numbered from the bottom to the top of the component (e.g., from the point of attachment of A1U1 on S to its tip). The diagram presented at the bottom of the figure details the geometric 3-D reconstruction of L in a scene, with respect to a coordinate system. The composite polygon (i.e., the set of four contiguous triangles) is a basic planar geometric model representing the shape and the size of L, where lL is the leaf midrib length and wL the leaf blade width. The position of L in space refers to its orientation and its position (coordinates of the point of attachment of L to P), with respect to the coordinate system. The leaf midrib direction is given by three angles: q, a rotation around the z-axis; j, a rotation around the y-axis; and y, a rotation around the x-axis. Definitions of symbols are given in Table 1.
where i = 1, , nM is the M number. Additionally, the curved shape of A1U1 is described by the evolution of the elevation angle (j) of every consecutive 20-cm-long segment (Se) along A1U1 (from plagiotropic Se at the bottom to orthotropic ones at the top of A1U1) as a function of lA1U1: jSei = f (lA1U1) (9)
where i = 1, , nSe is the Se number and nSe is the total number of Se along A1U1. From Equation 9, each consecutive I on A1U1 takes a j value (jI) given by the j value of its corresponding Se (jSe), as: jI i = jSeki Se1 , ki = 1 = 3 1 rand jSeki 1 ; jSeki 1 + jSenSe , ki = 2 , ..., nSe 4 4 (10)
With the A1U1 numbered arbitrarily, shoot diameter values (d10 A1U1) are calculated from shoot section area at 10-cm height according to a relative shoot rank (i/nA1U1) as:
where i = 1, , nM is the M number, ki = 1 l I j 20 + 1 is the corresponding Se number, x is the integer part of x, and rand[a, b] is a random number between a and b. Finally, A1U1 takes place in the scene after assigning qA1U1 a random number between 0 and 360.
CANOPY ARCHITECTURE OF COPPICE POPLAR
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May to September plant architecture reconstruction From the initial geometrical plant architecture described above, and for a selected date (e.g., May), lA1U2 is calculated for each A1U1 forming the plant as: lA1U2 = f (lA1U1) (11)
At last, the A1U1 reconstruction is completed by describing the foliage distribution along each A1U2 and A2U2, at the leaf scale. For each of them, total and individual leaf area values (AL and aL, respectively) are calculated according to lU2 and the relative M rank on U2 as: AL = f (l U2 ) (18)
To take into account potential variations in the size and shape of each A1U1 throughout the growing season (in terms of d increments and jSe variations), d10 A1U1 is recalculated as: d10 A1U1 = f (lA1U1 + lA1U2 ) (12)
i aL i = aL f nM
(19)
and then d b I, d t I and jI are recomputed from Equations 7, 8 and 10, respectively. At the A2U2 scale, the modeling process calculates the l of every A2U2 carried by each branched M on A1U1 as: i l I 1 j lA2U2 i = f lA1U1
where aL is the mean individual leaf area (= AL/nM) and i = 1, , nM is the M number. At this last scale, the leaf rate along the entire U2 was arbitrary fixed to 1, and temporal leaf abscission on U2 was not taken into account. For each individual leaf, lL, wL and petiole length (lP) values are calculated as: l L = f ( aL) wL = f (l L) l P = f (l L) (20) (21) (22)
(13)
where i = 1, , nM 3 is the M number, and nM 3 is explained by the presence of two consecutive vegetative buds and the A1U2 connected to the last three M at the top of A1U1. At this scale, the branching rate along the entire branched portion of A1U1 is arbitrary fixed to 1. Following Equations 5 and 6, A1U2 and A2U2 are, in their turn, divided into consecutive I units, described by their l according to a relative M rank on A1U2 or A2U2. In contrast with the geometrical description of A1U1, A1U2 and A2U2 are represented in the vegetative scene as conic frustums of height l (Equations 11 and 13, respectively), with d b and d t values given by: d b A1U2 = d t A1U1 d t A1U2 = f ( d b A1U2) d b A2U2 = f (lA2U2) d t A2U2 = f ( d b A2U2) (14) (15) (16) (17)
The final position and orientation in the scene of L and P are obtained after calculating q, a, j and twist (y) angle values as: rand[0, 360 ], i = 1 qPi = qPi -1 + dP, i = 2 , K , nM qL = qP i aPi = f nM i jL i = f nM i yL i = f nM (23)
(24)
(25)
The final position and orientation of A1U2 and A2U2 are obtained after calculating q, j and a (insertion angle) values as: qA1U2 = qA1U1 jA1U2 = rand[jSenSe , 90 ] rand[0, 360 ], i = 1 qA2U2 i = qA2U2 i -1 + dA2U2 , i = 2 , K , nM aA2U2 = c where d is the A2U2 divergence angle and c is a constant.
where i = 1, , nM is the M number and d is the P divergence angle. Finally, the plant structure is generated at metamer level from d b S and Equations 125, parameterized from field measurements for a given clone and date. Model outputs are a collection of geometric objects. Biometric measurements Equations 225 were parameterized with multi-scale biometric sampling. During the 2001 growing season, as the two poplar clones had only leafy A1U2 and A2U2 directly connected to A1U1, plant topology and geometry were assessed (i) at the clone scale, (ii) at stool and A1U1 scales in March (before bud
1158
burst), and (iii) at U2 (A1U2 and A2U2) and leaf (P and L) scales from May to September. Initial plant architecture parameterization (Equations 210) In mid-March, the d b of the 36 stools were measured above the ground with a caliper (dialMax-5921, Swiss Precision, Ashton, Switzerland) and the cumulated basal area was calculated. Additionally, nA1U1 was assessed (Equation 2). The cumulated basal area was divided into 10 (for baseline assessments) or five (for detailed assessments) equal classes (Cl). For each class, the mean basal stool diameter (d b S) was calculated and a stool with a d b close to the mean was selected and labeled from the 36 assessed stools. From these 15 selected and labeled stools, at the A1U1 scale, d10 (Equations 3, 4, 7 and 8) and l (Equations 49 and 1113) were measured on each alive A1U1 with a tape and the caliper, respectively. Additionally, on S labeled for detailed assessments, q was measured with a compass (KB-14/360 R, Suunto, Finland) for each A1U1. Using the same selection method as above, 45 A1U1 were selected (30 for baseline and 15 for detailed assessments) and labeled from three size classes (1, small; 2, medium; and 3, large) spanning the d10 A1U1 distribution range of all labeled stools. Finally, nM was measured along each A1U1 connected on an S labeled for detailed assessments (Equations 58), whereas lI (Equations 68) and d t (Equations 7 and 8) were measured only along labeled A1U1. May to September plant architecture parameterization (Equations 1125) Following the same multi-scale approach as above, measurements were carried out monthly from May to September (during the first and last 12 days of each month for Clones Tri and Gho, respectively). At the A1 scale, d10 A1U1 (Equation 12), d t A1U1 and d t A1U2 (Equations 14 and 15) were measured on all A1 connected on S labeled for detailed assessments. Additionally, the j value was measured for every consecutive Se along A1U1 in May, July and September (Equations 9 and 10). At the U2 scale, lA1U2 (Equations 11, 12 and 18) and lA2U2 (Equations 13, 16 and 18), nM (Equation 58) and nL (the number of leaves) of every U2 were measured along each labeled A1U1. At the A2U2 scale q, a, d b , d t (Equations 16 and 17, respectively), and lI (Equations 68) were measured on one selected and labeled A2U2 in three along each labeled detailed assessmentsA1U1. Finally, at the leaf scale, lL (Equations 2022), lP (Equation 22), qP, aP (Equation 23), jL (Equation 24) and yL (Equation 25) were measured at each leafy M on labeled U2 from May to September. Destructive sampling for leaf allometry From the unselected experimental poplar plots, a set of leaves was harvested separately for each clone during the entire growing season to parameterize Equations 1, 20 and 21. Leaves were sampled on A1U2 and A2U2 within the canopies and lL, wL and aL were measured in the laboratory with a laser area meter (CI-203, CID, Camas, WA). Hemispherical photographs The hemispherical photography method was used routinely to
estimate seasonal dynamics of canopy openness. Photographs were taken during uniform overcast sky conditions with a fish-eye lens (Nikon AF Fisheye 16 mm f/2.8D, Tokyo, Japan) mounted on a manually operated camera (Nikon FM2, Tokyo, Japan). A total of 15 below-canopy pictures were made 15 cm above the soil surface (i.e., top of the hemispherical lens) every week between May 1 and November 11. Five photographs were taken along the central row of the 36 assessed stools, and 10 from its two adjacent inter-rows.
Empirical functions and random deviation From the biometric measurements, empirical allometric relationships were described with polynomial, asymmetric sigmoid or power functions. The regression model was chosen based on the experimental data distribution, the best-fit value of the correlation coefficient of the adjustment (r 2) and a satisfactory random distribution of the residues (i.e., pass the SIGN test and/or the RUNS test). However, as an example, poplar trees are recognized to have a phyllotaxy angle (A2U2 angle around the A1) with a divergence ratio of 0.4. But, if dA2U2 is constant (i.e., 144), a top-down view of the A1U1 will show only the last five A2U2 at its tip. To be more realistic, a random deviation w was applied to selected empirical relationships (Equations 2, 3, 11, 13 and 2225) and constants (dA2U2, aA2U2, dP and aP), and was calculated as: w = rand[ -i , i] (26)
where i is the standard deviation of a mean or the maximal or the mean standard error of a constant pattern or of an empirical function, respectively. Hemispherical computer-generated images Plant images were computer-generated from the model outputs with the POV-Ray ray-tracing software program (Persistent of Vision Raytracer, Version 3.5). Each object or element of the scene was scaled to the appropriate geometric dimensions (e.g., height l, base radius d b /2 and top radius d t /2 for a conic frustum). The object was then rotated and translated according to its orientation and location in the scene (Figure 1). A specific texture was finally given to each object, describing its color. The resulting virtual coppice crop could then be looked at from any point of view, after having placed a virtual camera and light source in the scene. The light source (i.e., single and defined as direct sunbeams) was located a large distance from the scene at the zenith. Simulated coppice crop and hemispherical images were created with a virtual perspective camera with a viewing angle of 80 and 180, respectively. Following the experimental hemispherical photography method, five virtual hemispherical images were simulated along the central row of the 36 stools and 10 from the two adjacent inter-rows. Hemispherical photographs and virtual hemispherical images were processed with HemiView (Delta-T Devices, Cambridge, U.K.), a computer program for analyzing hemispherical images. For both kinds of image, the sky-map was constructed by dividing the sky into an array of sky annuluses
Table 3. Empirical functions at the stool (S) and shoot (A1U1) scales for Clones Ghoy (Gho) and Trichobel (Tri) for March and for the entire growing season of 2001. The relationships between Y and X were described with two regression models: (1) y = C0 + C1 x + C2 x 2 + C3 x 3 + C4 x 4 + C5 x 5 + C6 x 6; and (2) y = C 0 /(1 + (x / C 1)C2 ). Definitions of symbols are given in Table 1. X [min, max] Cl Model C0 C1 C2 C3 C4 C5 C6 n r2
Number
Month
[min, max]
Clone Gho 2 3 [2, 7] [0, 1] [0.1, 1.7] [10, 220] [0, 1] 1 1 0.535 2.135 3.275 1.675 2 1 369.4 7.636 1.262 0.280 1.516 3.067E3 1 0.061 0.325 1.023 1.367 0.672 1 3.809 0.428
[12, 47]
36 365 365 139 269
0.769 0.844 0.938 0.949 0.401
[0, 0.1]
4 5
3 3
[10, 220] [11, 58]
nA1U1 a10 A1U1i a bS lA1U1 nM lI i lA1U1
[0, 0.08]
d bS i nA1U1 d10A1U1 lA1U1 i nM
9 lA1U1 lA1U1 lA1 [5, 10] [0, 1] [0.2, 2.8] [28, 305] [0, 1] 1 1 0.532 2 1 456.3 7.593 1.558 0.172 1 0.076 1 22.15 0.611 1.360 1.25E4 2.082 3.184 [10, 255] [10, 220]
3 3 3
[0, 0.06] [0, 0.04] [16, 114]
[0, 1] [0, 1] [10, 220]
544 425 722 722 815
0.374 0.624 0.924
6, 9
[5, 90]
12
59
jSe1 jSenSe jSe1 jSenSe d10A1U1 2.152
[0.1, 2.9]
2 3
1 1 1 1 1 1 1
33.00 90.00 16.00 90.00
0.671 0.543 0.148 0.171 0.010
4.983 4.507 1.43E3 1.78E3 5.27E5
16.24 15.89 1.89E7
26.44 26.97
21.31 22.01 3.140 1.628 1.641
6.803 6.968
Clone Tri 2
[13, 30]
36 312 312 110 476
0.000 0.821 0.978 0.965 0.453
[0, 0.2]
4 5
3 3
[28, 305] [12, 50]
nA1U1 a10 A1U1i a bS lA1U1 nM lI i lA1U1
[0, 0.08]
d bS i nA1U1 d10A1U1 lA1U1 i nM
TREE PHYSIOLOGY ONLINE at http://heronpublishing.com lA1U1 lA1U1 lA1 [28, 305] [28, 305] [0, 1] [0, 1] [28, 305] 2 3 20.00 90.00 10.00 90.00 1 1 1 1 1 1 1 0.520 0.395 3.90E2 5.50E2 0.007 3.719 3.305 6.85E3 8.50E4 8.02E6 12.26 12.54 3.32E8
3 3 3
[0, 0.05] [0, 0.03] [6, 116]
650 465 747 737 652
0.550 0.686 0.964
6, 9
[5, 90]
12
59
jSe1 jSenSe jSe1 jSenSe d10A1U1
[0.2, 4.4]
20.13 22.89
16.26 19.90
5.184 6.635
1159
1160
Figure 2. Examples of empirical allometric relationships obtained from field measurements: the shoot /stool section area ratio (a10A1U1i /abS) versus its relative shoot rank on S (i/nA1U1) (Equation 3, Table 3); the relative internode length (lIi /lAIU1) versus its relative metamer rank on A1U1 (i/nM) (Equation 6, Table 3); and the elevation angle value of every consecutive 20-cm long segment along A1U1 (jSei) versus the shoot length (lA1U1) (Equation 9, Table 3) for poplar Clones Ghoy (Gho, left column) and Trichobel (Tri, right column). 59 are month numbers (MaySeptember, respectively) and Cl = 13 the A1U1 diameter class number (smalllarge, respectively). Definitions of symbols are given in Table 1.
defined by six zenith divisions (i.e., six annuli with 15 intervals).
Results Clonal differences in model parameterization In March 2001, d b S of Clone Gho was around 1.6 times smaller than that of Clone Tri but with a similar mean number of connected alive A1U1 (25 9 for Clone Gho and 22 6 for Clone Tri). Compared with Clone Gho, a weak variation on nA1U1 versus d b S (Empirical function 2, Table 3) and larger data ranges of d10 A1U1 (Empirical function 3, Table 3 and Figure 2) and lA1U1 (Empirical function 4, Table 3) were found for Clone Tri. At the I level, however, the two clones had a similar data range of nM as a function of lA1U1 (Empirical function 5, Table 3). Additionally, taking into account the shoot diameter class, empirical relationships obtained between the relative lI and its corresponding relative M rank on A1U1 (Empirical function 6, Table 3) showed similar data
ranges and data distribution shapes (i.e., trapezoidal sequential patterns) for the two clones (Figure 2). Internodes near the proximal and distal ends of every A1U1 were relatively short, but longer in their middle part (Figure 2), despite the presence of a weak depletion related to unfavorable weather at the beginning of summer in 2000 (data not shown). Finally, both clones had a large range in the curved shape of A1U1 according to its length (from plagiotropic to orthotropic shapes for the shortest to longest A1U1) (Empirical function 9, Table 3 and Figure 2). But no correlation between the curved shape of A1U1 and its position around the stool (in terms of location and orientation, according to the double-row design of the experimental plot) was found. The growing season of Clone Tri began in early April, about 26 days before that of Clone Gho. At the A1U1 scale, jSe values of the shortest A1U1, which were between the rows, decreased substantially during the first 2 months of the growing season from May to September (Empirical function 9, Table 3 and Figure 2). Additionally, for both clones, a common submodel was used to describe diameter evolutions along every
1161
Table 4. Empirical functions at the current-year growth-unit scale (U2) for Clones Ghoy (Gho) and Trichobel (Tri) for the entire growing season of 2001. The relationships between Y and X were described with three regression models: (1) y = C0 + C1x + C2 x 2; (2) y = C 0 /(1 + (x / C 1)C2 ); and (3) y = C 0 xC1 + C 2 . Definitions of symbols are given in Table 1. Scale Number Month Y [min, max] X [min, max] Cl Model C0 C1 C2 n r2
Clone Gho A1U2 11
A1U2 5 A2U2 13
5 6 7 89 59 59 59 5 6 7 89 59 59 5 6 7 8 9 59 59 5 6 79 89 59 59
lI
nM lA2U2i
[0.2, 10] [0.2, 19] [0.2, 35] [0.2, 50] [6, 31] [0.2, 7]
A2U2 16 U2 15, 17 Clone Tri A1U2 11
[0.2, 11] [0.2, 11] [0.2, 20] [0.2, 31] [0.2, 35] d b A2U2 [0.2, 0.5] [0.1, 0.2] d tU2 lI [0.2, 9] [0.2, 34] [0.2, 72] [0.2, 101] [0.2, 145] [3, 32] [0.2, 2]
lA1U1 [54, 185] [54, 185] [54, 185] [54, 185] lA1U2 [0.2, 50] i 1 l I j [0.6, 1] lA1U1 [0.5, 1] [0.4, 1] [0.4, 1] [0.4, 1] [0.4, 1] lA2U2 [0.2, 45] d bU2 [0.2, 0.5] lA1U1 [72, 301] [72, 301] [72, 301] [72, 301] [72, 301] lA1U2 [0.2, 145] i 1 l I j [0.2, 1] lA1U1 [0.1, 1] [0.1, 1] [0.1, 1] [0.1, 1] [0.1, 1] [0.1, 1] [0.1, 1] lA2U2 [0.2, 73] d bU2 [0.2, 0.9]
13 13 13 13 13 1 2 3 3 3 3 13 13 13 13 13 13 13 13 1 2 3 2 3 2 3 3 13 13
2 2 2 2 3 1 1 1 1 1 1 3 1 3 3 3 3 3 3 3 3 3 3 3 3 3 3 1 1
8.892 16.89 31.69 41.87 5.173 2.508 14.62 19.58 18.56 18.61 15.04 0.178 3.13E3 1.30E2 4.41E3 1.20E3 2.73E4 1.936 1.357 4.570 11.49 25.83 38.56 38.53 67.78 75.86 0.263
92.53 79.11 86.49 87.57 0.442 4.980 42.13 64.70 57.15 54.05 39.43 0.261 0.370 1.385 1.384 1.700 1.987 2.296 0.563 2.281 3.389 1.942 7.360 2.784 9.258 3.911 4.232 7.90E3 0.346
2.547 3.966 4.360 3.977 22.410 39.34 26.96 18.87 4.530 0.171 0.306 0.155 0.469 1.294 1.559
43 41 40 83 189 211 875 214 215 191 376 355 372 40 46 34 39 46 319 610 440 362 457 353 1105 404 767 171 298
0.738 0.813 0.819 0.767 0.962 0.061 0.146 0.383 0.415 0.362 0.374 0.811 0.338 0.738 0.813 0.819 0.767 0.767 0.962 0.061 0.146 0.383 0.415 0.362 0.374 0.362 0.362 0.811 0.338
A1U2 5 A2U2 13
nM lA2U2i
A2U2 16 U2 15, 17
[0.2, 7] [0.2, 13] [0.2, 26] [0.2, 40] [0.2, 42] [0.2, 73] [0.2, 84] d bA2U2 [0.2, 0.9] [0.1, 0.3] d tU2
A1U1 (d I i = d10 A1U1(a 1 l I j lA1U1 + b), where i = 1, , nM is the M number, a = 0.871 0.008 and b = 1.029 0.004; n = 156, r 2 = 0.998) (see Equations 7 and 8), despite the clones having different A1 size ranges (Empirical function 12, Table 3). At the U2 and leaf scales (Tables 4 and 5 and Figure 3), strong contrasts were found between the two clones in the empirical allometric relationships observed. For both clones, lA1U2 and lA2U2 were always positively correlated to the shoot diameter class (Empirical functions 11 and 13, Table 4 and Figure 3). But, in contrast with Clone Tri, Clone Gho showed a relatively constant lA1U2 for the largest A1U1 diameter class, highlighted by an asymmetric sigmoid function fit between lA1U2 and lA1U1, compared with a power function for Clone Tri (Empirical function 11, Table 4 and Figi
ure 3). Similar regression models were used to describe the acropetal increase of lA2U2 on A1U1 for both clones (Empirical function 13, Table 4 and Figure 3), although stronger data variability was observed for Clone Gho (Figure 3). Additionally, Clone Gho was characterized by the presence of unbranched M along the first half of its A1U1 (Empirical function 13, Table 4 and Figure 3). However, both clones exhibited a branching rate of about 1 with common dA2U2 values (141 13, n = 352) along the entire branched portion of every A1U1. Finally, aA2U2 had a range of 40 5 for Clone Gho and 49 11 for Clone Tri. Whereas a single relationship described nM versus lA1U2 for the entire growing season, and for all A1U1 diameter classes (Empirical function 5, Table 4), the description of A1U2 and A2U2 entities (i.e., I) required relationships for
1162 Table 5. Empirical functions at the leaf scale (petiole, P and leaf lamina, L levels) for clones Ghoy (Gho) and Trichobel (Tri) for the entire growing season of 2001. The relationships between Y and X were described with two regression models: (1) y = C0 + C1x + C2 x 2 + C3 x 3 + C4 x 4 + C5 x 5 + C6 x 6; and (2) y = C 0 /(1 + (x / C 1)C2 ). Definitions of symbols are given in Table 1. 0.968 0.935 0.500 0.774 0.751 0.302
CASELLA AND SINOQUET 0.990 0.997 0.080 0.696 0.502 0.090
each A1U1 diameter class and observation date. Compared with the description of A1U1, lI and nM data ranges observed at the U2 scale were less comparable between the clones. Relative lI ranges were smaller for Clone Gho than Clone Tri, suggesting a higher leaf initiation rate for Clone Gho. Finally, at the leaf scale (Table 5), except for the distribution of jL along U2 (Empirical relationship 24, Table 5), A1U2 and A2U2 characteristics (Table 5) were described with a single relationship for each clone. For both clones, AL of A1U2 was larger than that of A2U2 and increased with increasing lA1U2 or lA2U2 (Empirical function 18, Table 5). The distribution of aL along A1U2 (Figure 3) or A2U2 (Empirical function 19, Table 5) showed similar data ranges and sequential patterns between the clones with the shape of the relationship progressing from parabolic in May to trapezoidal in September. Additionally, the two clones exhibited a common dP value (141 13, n = 352) along the entire leafy part of every U2. Compared with Clone Tri, leaves of Clone Gho were different in many geometric characteristics: heart-shaped leaf blades (wL = 0.809lL, r 2 = 0.865; lL = 2.043 aL, r 2 = 0.912; and aL = 0.593lLwL, r 2 = 0.988; n = 658) compared with lanceolate ones for Clone Tri (wL = 0.582lL; l L = 2.565 aL , r 2 = 0.976; and aL = 0.631lLwL, r 2 = 0.996; n = 821), aL was about four times smaller, lP was 75% of lL versus 30% for Clone Tri, and larger data ranges described jL (Empirical function 24, Table 5) and yL, which had a range of 0 80 (n = 141) versus 0 20 (in May, n = 105) and 0 40 (July and September, n = 386) for Clone Tri. Evaluation of the virtual coppice crop reconstruction method Throughout the entire growing season of 2001, model outputs were compared with field measurements to test features ranging from the characteristics of individual components of a plant (Figure 4, Table 6) to the more complex structure of a canopy (Figures 57). From May to September and for both clones, results reported in Figure 4 and Table 6 show that the modeling process recreates the geometry of a plant (see also Figure 1). First, the seasonal time course of the simulated total number of potential leafy M per A1 (i.e., M on U2 only) is strongly correlated with the measured one (Figure 4). Second, as observed from the experimental results, mean lA1U2 and lA2U2 derived from model outputs are positively correlated with the A1U1 diameter class (Table 6). Third, the clonal contrast in the acropetal increase of lA2U2 on A1U1 is well recreated by the modeling process (Table 6). Finally, at the leaf scale, spatial and temporal variations in AL are taken into account by the model (Figure 1 and Table 6). At the plot level, the quality of the spatial and temporal distributions of every plant entity in a scene is shown by the synthesized images in Figure 5. For each clone, the reconstruction method seems to recreate the geometry of the canopy in accordance with the double-row design of the experiment. Clonal and temporal differences in coppice structure (e.g., A1 geometry) and canopy morphology (e.g., L geometry), detailed in the preceding paragraph, are clearly recreated (Figure 5) as the
r2
7.76E8
C6
92 1311 5676 559 276 105 386
84 423 4126 386 154 141
2.93E3 C4 0.879 C3 C2 0.128 0.133 2.918 1.920 4.901 11.68 4.347 4.093 81.700 79.30
0.168 0.986 21.86
21.14 52.19 2.026 6.897
Model
71.48 6.310 99.90 55.87
C0
[min, max]
lA1U2 lA2U2 i/nA2U2 lL i/nM i/nM
[3, 48] [0, 35] [0, 1] [0, 7] [0, 1] [0, 1]
[min, max]
AL AL aL i / aL lP aPi jLi
Month
59 59 59 59 59 5, 6, 9
Clone Gho 18 18 19 22 23 24
Number
Clone Tri 18 18 19 22 23 24 24
59 59 59 59 59 5 6, 9
AL AL aL i / aL lP aPi jLi jLi
[200, 5132] [5, 1674] [0, 2.57] [0, 6] [10, 80] [85, 0] [70, 90]
[70, 613] [2, 265] [0, 2.68] [0, 6] [5, 120] [85, 80]
lA1U2 lA2U2 i/nA2U2 lL i/nM i/nM i/nM
[6, 144] [0, 80] [0, 1] [0, 22] [0, 1] [0, 1] [0, 1]
1 1 1 2 1 1 1
1 1 1 2 1 1
0.767 2.782 66.14 52 22 23.72
29.12
C1
0.107 4.314 3.027 4.496 443.400
1002.000
601.20
2.43E5
C5
1163
Figure 3. Examples of empirical allometric relationships obtained from field measurements: the shoot leader length (lA1U2) versus the shoot length (lA1U1); the branch length (lA2U2i, for the shoot diameter class 3 only) and the individual/mean individual U2 leaf blade area ratio (aL i / aL) versus its corresponding relative metamer rank (i/nM) on A2U2 and A1U2, respectively, for poplar Clones Ghoy (Gho, left column) and Trichobel (Tri, right column). Numbers 59 indicate month (MaySeptember, respectively). Definitions of symbols are given in Table 1.
Figure 4. Measured and simulated total number of leafy metamers, at the shoot scale, over the 2001 growing season ( = May, = June, = July, = August and = September). Cl = 13 is the shoot diameter class number (smalllarge, respectively). Relationships were fitted to the linear regression y = ax + b. Dotted lines depict the confidence interval at P < 0.05.
images show the strong contrast in growth potential expressed by the two clones. To assess the spatial aL distribution, virtual hemispherical pictures were compared with real hemispherical photographs (Figures 6 and 7, respectively). A visual inspection (Figure 6) shows that A1, A2, leaf and canopy gap sizes, shapes and spatial distributions are correctly recreated
throughout the growing season for the two clones (Figure 6), implying that the modeling process recreates the spatial and temporal aL distribution within the contrasting canopies. However, due to the restricted dimensions of the experimental plots and the surrounding forest landscapes, a dense border was observed on the horizon of the real hemispherical photo-
1164
Table 6. Basic structural plant and canopy parameters derived from the 3-D CPCA model outputs describing the poplar Clones Ghoy (Gho) and Trichobel (Tri). Results (MaySeptember) were generated from a population of 36 virtual plants by the modeling operation, linearly corrected between two consecutive months ( 6 days, taking into account the delay of 12 days between the assessment of the two clones), and displayed by shoot diameter class (Cl), relative branch position on A1U1 (l I lA1U1) or within the canopies. Abbreviations: n = number of; l = length (cm); DlI = change in lI between May and September, the internode elongation growth potential; DnL = change in nL between May and September, the leaf initiation rate; and aL = leaf blade area (cm 2), characterizing an average shoot (A1U1), shoot leader (A1U2), branch (A2U2), metamer (M), internode (I) or leaf blade (L) of an individual or a population of plants. Definitions of symbols are given in Table 1. Bars over symbols denote means. Plant
A1U1 Cl n l nI lI
A1U2 l nI DlI 2.4 aL
A2U2
lI lA1U1
0.751 0.50.75 00.5 0.751 0.50.75 00.5 0.751 0.50.75 00.5 0.41 0.751 0.50.75 00.5 0.751 0.50.75 00.5 0.751 0.50.75 00.5 01
nI
DlI 1.5 0.8 0.8 0.5 0.4 0.0 0.5 0.1 0.5 4.8 3.9 0.7 3.7 2.0 0.4 2.0 0.2 0.1 1.9
aL
Clone Gho 3 3
171
46
3.7
5.441.3
1126
9.319
120
36
3.3
4.333.6
1024
2.1
9.516
16
63
23
2.7
1.911.1
613
1.3
12.412
13
25
89 279
29 45
3.1 6.2
2.919.9 12.4126.7
817 729
1.9 5.2
11.014 39.3160
11 9 4 8 8 2 5 4 0 13 10 9 24 8 7 18 4 4 6 21
5.718.9 4.79.7 1.22.0 3.85.4 1.72.5 0.10.1 1.41.9 0.30.5 1.73.2 13.752.0 6.314.1 1.02.4 6.224.5 1.93.9 0.31.0 1.83.8 0.70.9 0.20.3 1.65.3
817 713 56 710 57 11 56 22 69 715 68 24 49 34 13 34 23 11 35
7.910 6.98 3.24 5.66 3.94 1.42 3.13 2.23 5.16 40.166 30.038 19.028 37.445 24.236 11.418 25.729 14.917 7.89 2632
Clone Tri 3 2
184
34
5.4
7.049.9
517
3.6
32.983
15
86
21
4.1
2.510.6
26
2.0
28.940
13
22
124
26
4.8
4.429.7
310
3.6
32.285
Canopy A1U2 Cl Clone Gho 3 2 1 13 Clone Tri 3 2 1 13 n nM DnL 16 14 7 9 22 12 4 7 A2U2 Layer depth (m) n nM DnL 15 7 2 3 16 6 1 2
111 200 586 897 78 148 554 780
11932914 19314776 36157532 673815221 5502243 7292468 13813547 26598258
1.62.6 1.01.6 01.0 02.6 2.54.4 1.62.5 01.6 04.4
353 2079 9231 11663 300 1363 14603 16267
30398308 1726531135 4877466958 69078106400 25367327 744015144 3356551745 4354174217
graphs (Figure 6). Taking this observation into account, the seasonal time course of the spatial aL distribution within each canopy was assessed by comparing canopy openness values calculated from the virtual and real hemispherical images for each sky-map zenith division (Figure 7). A strong direct correlation was found between simulated and measured canopy
openness values for each clone for the first four zenith angles (between 0 and 60). But the border effect (exceptionally strong between 75 and 95) was not recreated by the model because virtual crop canopies were limited to 100 plants. By integrating these values over a restricted hemispherical viewing angle of 120, temporal courses of the measured and simulated
1165
Figure 5. Examples of virtual 2-year-old poplar crop images, computer-generated from the 3-D CPCA model outputs by the POV-Ray software for poplar Clones Ghoy (Gho) and Trichobel (Tri) in May and September 2001. Crop canopies, consisting of 100 plants each, are viewed from the South to the North. Axis lengths represent 1 m.
Figure 6. Examples of visual comparisons between fish-eye photographs (top line) and fish-eye images (bottom line) generated from the 3-D CPCA model outputs by the POV-Ray software for poplar clones Ghoy (Gho; ad, between-row location) and Trichobel (Tri; eh, inside-row location). Crop canopies consist of 100 plants each.
canopy openness values could be compared over the entire growing season for each clone (Figure 7). The modeling process recreates (i) weak row-related canopy closure dynamics
and the strong heterogeneity observed throughout the entire growing season for the canopy of Clone Gho (Figures 57), and (ii) the fast (less than 3 months) canopy closure of Clone
1166
Figure 7. Measured (mes) and simulated (sim) canopy openness over six sky-map zenith divisions ( , = 015; , = 1530; , = 3045; , = 4560; , = 6075; and , = 7590) over the 2001 growing season, and temporal courses of canopy openness (lines = measured and symbols = simulated) for poplar Clones Ghoy (Gho, left column) and Trichobel (Tri, right column). Open and closed symbols show between- and inside-row locations, respectively. Vertical and horizontal bars indicate the standard deviation of the mean (n = 5 or 10 for the inside- or between-row locations, respectively). Letters indicate significant differences (t-test, P < 0.05) between row locations and results obtained from field measurements and the modeling process. Relationships between simulated and measured canopy openness were fitted, for four zenith angles (060), to the linear regression y = ax + b. Dotted lines depict the confidence interval at P < 0.05.
Tri, which was constant (at about 90% compared to a maximum of 60% for Clone Gho) after mid-June (Figures 57). Discussion Clonal characteristics A considerable number of studies have shown that stands of certain highly productive poplars (often P. trichocarpa and its hybrids) quickly achieve high LAI values (up to 10 or more), whereas less productive ones (often Euramerican hybrids) may reach only one-third to one-half of those values (DeBell et al. 1996, Heilman et al. 1996, Casella and Ceulemans 2002). But this genetic tendency is not always apparent because the development of plant LAI is often under strong environmental control (site climate, plantsoilnutrient interactions, pests and diseases) (Liu and Dickmann 1992a, 1992b, Stettler et al. 1996, Ceulemans and Deraedt 1999, Dickmann et al. 2001). Because leaf physiological processes (i.e., maximal photosynthetic electron transport capacity and carboxylation rate) seem to be relatively constant over the Populus spp. (Niinemets et al. 1998, Casella and Ceulemans 2002, Ripullone et al. 2003), canopy LAI development and leaf angular and spatial distributions over a growing season seem to be the most pronounced genetic traits expressing the large range in yield of this species. Thus, under the same environmental conditions, these clonal contrasts could be explained through the expression of contrasting genotypic traits related to their potential to maximize
ei by an optimal coppice and canopy structure arrangement over the growing season (number of A1, A2 and L, and their geometry in space). The broad range of features detailed in this paper and generated by the 3-D CPCA model (Table 6) are now used to provide some explanations for the difference in performance of our two clones. Although both the empirical functions and the 3-D CPCA model do not consider the A1 mortality and L abscission in the vegetative period, potential LAI evaluated from model outputs could be used for a clonal comparison between May and July because the leaf longevity of most poplar clones is between 90 and 120 days (Zavitkovski 1981, Ceulemans et al. 1992) and canopy closure of Clone Tri occurred in late July (Figure 7). As expected, throughout the growing season of 2001, the clonal differences in coppice and canopy structures of our two clones resulted in striking differences in plant phenology, and A1, A2, and L morphology and growth characteristics. Compared with Clone Tri, the growing season for Clone Gho was shortest (about 5 months compared with 7 for Clone Tri), and began about 26 days later than for Clone Tri (Figure 7). In March, the two coppices showed a similar A1U1 density (Table 6) but, despite a similar number of I, the A1U1 of Clone Gho were on average 1.4 times shorter than those of Clone Tri (Table 6). In contrast with several results for various natural (e.g., P. trichocarpa) and hybrid (e.g., Euramerican poplars) poplar clones (Ceulemans et al. 1990, Dunlap et al. 1992, Ceulemans and Isebrands 1996), a small number of lateral
1167
sylleptic A2 characterized the A1U1 of our two clones (< 5%, and only on A1U1 from the large diameter class). From May to September, despite differences in coppice structure (in terms of the mean size of U2) and canopy morphology (in terms of the mean number and size of L) (Table 6), the two clones displayed similar vertical profiles for lA1U2, lA2U2 and aL. Mean lU2 and aL increased with distance along A1U1 (Table 6). These branch and foliar distribution patterns within a canopy, related to a vertical light profile (Figure 7), agree with previous experimental results describing the spatial distribution of A2 and L characteristics within poplar canopies (Ceulemans et al. 1990, Chen 1992, Casella and Ceulemans 2002). Because no sylleptic A2U2 had lengthened on A1U2 for both clones, and because Clone Gho was characterized by the persistence of vegetative buds along the first half of its A1U1 (Figures 1, 3 and Table 6), its coppice displayed a A2U2 density about 1.4 times lower than that of Clone Tri (Table 6). But, despite a later start to its growing season and shortest U2 (about 1.5 times on average, Figure 3 and Table 6), the potential number of leafy M characterizing the canopy of Clone Gho was about 1.5 times greater than for Clone Tri (Table 6). At the plant and canopy scales, these results suggest that Clone Gho had a higher mean leaf initiation rate (DnL) and a smaller internode elongation growth potential (DlI) throughout the growing season than Clone Tri (Table 6). However, dividing the canopy volume of each clone into three horizontal layers (Table 6), A1U2 and A2U2 DnL and DlI values varied strongly within the two canopies (Table 6). For both clones, DnL and DlI values were low near the bottom of the canopy and increased with height (Table 6). These patterns, showing a simultaneous increase in DnL and DlI in response to irradiance, agree with the results reported for Clone Robusta (P. deltoides P. nigra) by Pieters et al. (1999). But, if only the top part of the canopies was considered (Table 6), DnL of A1U2 was about 1.4 times lower in Clone Gho than in Clone Tri. Because, for both clones, the growth rate of an A1U2 connected on a large A1U1 (i.e., Cl = 3, Table 6) was not limited by PAR availability, this observation suggests that Clone Gho had a lower DnL than Clone Tri. This contrasting genotypic trait agrees with the results reported by Ceulemans et al. (1988), showing that leafy A1U1 of Clone Gho displayed DnL about 1.3 times lower than those of Clone Tri. However, because Clone Tri closed its canopy much earlier and more tightly than Clone Gho (Figure 7), the DnL values of Clone Gho were higher than those of Clone Tri at the lower parts of their canopies (Table 6). This agrees with the results reported for Fraxinus pennsylvanica Marsh. (Remphrey et al. 2002) and with the fact that, at the canopy scale, Clone Ghoy displayed a larger number of leafy M than Clone Tri (Table 6). Additionally, the comparison between their DlI values simulated at the A1U1 and A1U2 levels (Table 6A) suggested that I elongation rates of both clones were not optimal in the second growing season. These results corroborate the assertion that the increasing proportion of plant mass that does not participate in growth causes a continuous decline in growth rates of new U2 (Lord et al. 1993). Finally, at the leaf level, further genotypic traits strongly dif-
ferentiated our two clones in terms of L sizes, shapes and distribution along a U2. Compared with Clone Tri, U2 of Clone Gho were shortest and displayed a greater number of smallest L (Table 6). Additionally, in contrast with the relatively horizontal position of L of Clone Tri, those of Clone Gho were highly angled (both aL and yL, see results). As a result of the contrasting genotypic traits expressed by our two clones, we expect natural or hybrid poplar clones characterized by a low DnL, small and highly angled L, and thus lower DlI (characterizing the canopy of Clone Gho) to have a lower e i than those expressing high DnL with big and horizontal L (beneficial for an optimal DlI, as for Clone Tri). These hypotheses corroborate a large range of results and analyses concerning the effects of canopy LAI development and productivity (Heilman and Stettler 1985, DeBell et al. 1996, Heilman et al. 1996, Pontailler et al. 1999, Laureysens et al. 2000, Casella and Ceulemans 2002, Tubby and Armstrong 2002), the relationships between leaf morphology and shoot light capture (Takenaka 1994), and the contrasting temporal course of e i displayed by our two poplar canopies (Figure 7). Finally, all these results, based on a detailed set of field measurements, explain observed differences in crop-level performance, both in terms of LAI (1.7 versus 6.0 for Clones Gho and Tri, respectively, in July) and biomass production (4 versus 11 Mg DM ha 1 year 1 for Clones Gho and Tri, respectively) (authors unpublished results). Virtual coppice crop reconstruction method The present paper proposes a method to reconstruct the plant architecture of coppice poplar from an extensive set of allometric rules, derived from a hierarchical sampling in the studied canopies. Similar approaches have been proposed to study the space occupation by pine trees (Whitehead et al. 1990) and make 3-D plant mock-ups for botanical and landscape purposes (De Reffye et al. 1988, 1991, Barczi et al. 1997). The reconstruction methods allow one to generate plant mock-ups with the same topological and geometrical properties as the plant population for which the architectural rules have been established. Reconstruction methods usually do not deal with the reconstruction of a given plant in the population. Moreover, because reconstruction methods use a sequence of empirical relationships that show uncertainty, it is necessary to assess the quality of the resulting plant. In this paper, fish-eye images of the virtual canopies were compared to real photographs taken in the field. Comparing virtual and real images is a common way to assess the quality of 3-D plant mock-ups for space occupation. Previous comparisons have been qualitative (e.g., Sinoquet and Rivet 1997) or quantitative, i.e., the comparison of variables computed from the photographs (Ivanov et al. 1995, Rakocevic et al. 2000). The quantitative evaluation of 3-D plant mock-ups is more satisfactory, especially for reconstruction methods, for two reasons. First, reconstructed plants do not match a given real plant, as reconstruction rules are generic to all plants in the population for which the architectural rules have been established. Figure 6 provides an illustration of the mismatch, where the plants in the virtual canopy do not visually match the plants seen in the real photographs. Second,
1168
a quantitative assessment of reconstruction quality allows one to evaluate easily the effect of combining several empirical rules. For the first reason, such a criterion of reconstruction quality (e.g., canopy openness in this study) must be defined at the plant population scale, not the individual plant scale. The criterion can be used to refine the set of architectural rules and identify the main architectural parameters that determine the 3-D space occupation. In this study, we used a large set of reconstruction rules in order to increase the reconstruction quality (Figure 7). The next step would be to evaluate simplified reconstruction methods (i.e., by replacing some rules with assumptions, e.g., constant internode length), and then to define the minimum data sets to be measured in the field for a given reconstruction quality. Conclusion The 3-D CPCA model, based on a detailed set of empirical relationships obtained from field measurements, recreates the spatial aL distribution within contrasting SRWC poplar canopies. This study shows that, as suggested by Casella and Ceulemans (2002), foliage cannot be accurately described by assuming it to be randomly distributed within horizontal layers of Populus canopy. From a methodological point of view, the 3-D CPCA model is likely to find several applications. (i) It could be used to make sensitivity analyses of the reconstruction method to identify the relevant parameters and define the minimum data set to be measured in the field to feed the model. (ii) Because the 3-D CPCA model expresses the spatial geometry of stools, U2 and leaves of a population of plants, it could be used as an input to 3-D models dealing with the distribution of light interception (e.g., RATP, Sinoquet et al. 2001), rainfall interception (Gash 1979), wind speed (Daudet et al. 1999), photosynthesis and transpiration (e.g., RATP), and sap flow (Dauzat et al. 2001) activities within a canopy. With the accurate description of the aL distribution within our canopies, a further application of the 3-D CPCA model could be the exploration of the effects of foliage clumping on radiation interception. Error in radiation interception resulting from the assumption that foliage is randomly distributed could be estimated. (iii) The 3-D CPCA model could be used to test all these enumerated models at different scales (from branch to canopy) to propose comprehensive analysis of the relationships between leaf morphology and branchcanopy functions (e.g., light capture, manipulating shoot and branch inclination angles, petiole length, leaf shape, size and orientation) (Takenaka 1994) that will lead to increased productivity. Radiative transfer models could be used to predict the potential growth of individual poplar clones and ensure the efficient selection of those that will remain for the final crop. Finally, (iv) data sets obtained with canopies measured for several years could be used to test models of canopy architecture dynamics (Sinoquet and Le Roux 2000).
Acknowledgments
rie Curie Fellowship of the European Community program Quality of Life and Management of Living Resources under contract number QLK5-CT-2001-50583. The authors gratefully acknowledge R. Ceulemans (University of Antwerp, Belgium) for useful discussions and review comments, as well as M. Baldwin (Forest Research) for improving the English of this manuscript.
References Armstrong, A. 1997. The United Kingdom network of experiments on site/yield relationships for short rotation coppice. Note 294. Forestry Commission Research Information, Forestry Commission, Edinburgh, 6 p. Barczi, J.F., P. de Reffye and Y. Caraglio. 1997. Essai sur lidentification et la mise en uvre des paramtres ncessaires la simulation dune architecture vgtale: le logiciel AMAPsim. In Modlisation et Simulation de lArchitecture des Vgtaux. Eds. J. Bouchon, P. de Reffye and D. Barthlmy. INRA Editions, Paris, pp 205254. Cannell, M.G.R. 1989. Physiological basis of wood production: a review. Scand. J. For. Res. 4:459490. Cannell, M.G.R., L.J. Sheppard and R. Milne. 1988. Light use efficiency and woody biomass production of poplar and willow. Forestry 61:125136. Casella, E. and R. Ceulemans. 2002. Spatial distribution of leaf morphological and physiological characteristics in relation to local radiation regime within the canopies of 3-year-old Populus clones in coppice culture. Tree Physiol. 22:12771288. Chen, S.G. 1992. Modeling architecture and the light regime of poplar canopies using fractal approach. Ph.D. Thesis, Dept. Biology, Univ. Antwerp, Belgium, 200 p. Chen, S.G., R. Ceulemans and I. Impens. 1994a. A fractal-based Populus canopy structure model for the calculation of light interception. For. Ecol. Manage. 69:97110. Chen, S.G., B.Y. Shao, I. Impens and R. Ceulemans. 1994b. Effects of plant canopy structure on light interception and photosynthesis. J. Quant. Spectrosc. Radiat. Transf. 52:115123. Ceulemans, R. and W. Deraedt. 1999. Production physiology and growth potential of poplars under short-rotation forestry culture. For. Ecol. Manage. 121:923. Ceulemans, R. and J.G. Isebrands. 1996. Carbon acquisition and allocation. In Biology of Populus and its Implications for Management and Conservation. Eds. R.F. Stettler, H.D. Bradshaw, P.E. Heilman and T.M. Hinckley. NRC Research Press, Ottawa, Canada, pp 355399. Ceulemans, R., I. Impens and V. Steenackers. 1988. Genetic variation in aspects of leaf growth of Populus clones, using the leaf plastochron index. Can. J. For. Res. 18:10691077. Ceulemans, R., R.F. Stettler, T.M. Hinckley, J.G. Isebrands and P.E. Heilman. 1990. Crown architecture of Populus clones as determined by branch orientation and branch characteristics. Tree Physiol. 7:157167. Ceulemans, R., G. Scarascia-Mugnozza, B.M. Wiard, J.H. Braatne, T.M. Hinckley, R.F. Stettler, J.G. Isebrands and P.E. Heilman. 1992. Production physiology and morphology of Populus species and their hybrids grown under short rotation. I. Clonal comparisons of 4-year growth and phenology. Can. J. For. Res. 22:19371948. Ceulemans, R., A.J.S. McDonald and J.S. Pereira. 1996. A comparison among eucalypt, poplar and willow characteristics with particular reference to a coppice, growth-modeling approach. Biomass Bioenergy 11:215231.
This research was supported by a grant from Forest Research, an executive agency of the Forestry Commission, and by a Ma-
CANOPY ARCHITECTURE OF COPPICE POPLAR Daudet, F.A., X. Le Roux, H. Sinoquet and B. Adam. 1999. Wind speed and leaf boundary layer conductance variation within tree crown: consequences on leaf-to-atmosphere coupling and tree functions. Agric. For. Meteorol. 97:171185. Dauzat, J., B. Rapidel and A. Berger. 2001. Simulation of leaf transpiration and sap flow in virtual plants: model description and application to a coffee plantation in Costa Rica. Agric. For. Meteorol. 109: 143160. DeBell, D.S., G.W. Clendenen, C.A. Harrington and J.C. Zasada. 1996. Tree growth and stand development in short-rotation Populus plantings: 7-year results for two clones at three spacings. Biomass Bioenergy 4:253269. Deraedt, W. and R. Ceulemans. 1998. Leaf phenology in the first and second year of growth in a coppice biomass plantation. Biomass for Energy and Industry, CEC, Carmen, Germany, pp 969971. De Reffye, P., C. Edelin, J. Franon, M. Jaegger and C. Puech. 1988. Plant models faithful to botanical structure and development. Comput. Graph. 22:151158. De Reffye, P., E. Elguero and E. Costes. 1991. Growth units construction in trees: a stochastic approach. Acta Biotheor. 39:325342. Dickmann, D.I., J.G. Isebrands, J.E. Eckenwalder and J. Richardson. 2001. Poplar culture in North America. NRC Research Press, Ottawa, Canada, 397 p. Dunlap, J.M., P.E. Heilman and R.F. Stettler. 1992. Genetic variation and productivity of Populus trichocarpa and its hybrids. V. The influence of ramet position on 3-year growth variables. Can. J. For. Res. 22:849857. Eckenwalder, J.E. 2001. Descriptions of clonal characteristics. In Poplar Culture in North America. Eds. D.I. Dickmann, J.G. Isebrands, J.E. Eckenwalder and J. Richardson. NRC Research Press, Ottawa, Canada, pp 331382. Gash, J.H.C. 1979. An analytical model of rainfall interception by forests. Quart. J. R. Met. Soc. 105:4355. Godin, C., E. Costes and H. Sinoquet. 1999. A method for describing plant architecture which integrates topology and geometry. Ann. Bot. 84:343357. Grassi, G., G. Gosse and G. Dos Santos. 1990. Biomass for energy and industry. Vol. 1. Policy, environment, production and harvesting. Elsevier Applied Science, London, 697 p. Hall, F., R.A.A. Oldeman and P.B. Tomlinson. 1978. Tropical trees and forests. An architectural analysis. Springer-Verlag, Berlin, 411 p. Heilman, P.E. and R.F. Stettler. 1985. Genetic variation and productivity of Populus trichocarpa and its hybrids. II. Biomass production in a 4-year plantation. Can. J. For. Res. 15:384388. Heilman, P.E., T.M. Hinckley, D.A. Roberts and R. Ceulemans. 1996. Production physiology. In Biology of Populus and its implications for management and conservation. Eds. R.F. Stettler, H.D. Bradshaw, P.E. Heilman and T.M. Hinckley. NRC Research Press, Ottawa, Canada, 539 p. Ivanov, N., P. Poissard, M. Chapron and B. Andrieu. 1995. Computer stereo plotting for 3-D reconstruction of a maize canopy. Agric. For. Meteorol. 75:85102. Laureysens, I., E. Casella and R. Ceulemans. 2000. Productivity, basal area and shoot density of different poplar clones in a coppice culture. In Topics in Ecology: Structure and Function in Plants and Ecosystems. Eds. R. Ceuleman, J. Bogaert, G. Deckmyn and I. Nijs. Univ. Antwerp, Wilrijk, Belgium, pp 2938. Law, B.E., A. Cescatti and D.D. Baldocchi. 2001. Leaf area distribution and radiative transfer in open-canopy forests: implications for mass energy exchange. Tree Physiol. 21:777787.
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Liu, Z. and D.I. Dickmann. 1992a. Responses of two hybrid Populus clones to flooding, drought and nitrogen availability. I. Morphology and growth. Can. J. Bot. 70:22652270. Liu, Z. and D.I. Dickmann. 1992b. Responses of two hybrid Populus clones to flooding, drought and nitrogen availability. II. Gas exchange and water relations. Can. J. Bot. 71:927938. Lord, D., S. Morissette and J. Allaire. 1993. Influence of light intensity, night air temperature and CO2 concentration on growth of containerised black spruce seedlings grown in the greenhouse. Can. J. For. Res. 23:101110. Macpherson, G. 1995. Home-grown energy from short-rotation coppice. Farming Press, Ipswich, U.K., 214 p. Milne, R., M. Sattin, J.D. Deans, P.G. Jarvis and M.G.R. Cannell. 1992. The biomass production of three poplar clones in relation to intercepted solar radiation. For. Ecol. Manage. 55:114. Niinemets, ., O. Kull and J.D. Tenhunen. 1998. An analysis of light effects on foliar morphology, physiology, and light interception in temperate deciduous woody species of contrasting shade tolerance. Tree Physiol. 18:681696. Pieters, G.A., M.E. van den Noort and J.A. van Nijkerken. 1999. Growth adaptation of leaves and internodes of poplar to irradiance, day length and temperature. Tree Physiol. 19:933942. Pontailler, J.Y., R. Ceulemans and J. Guittet. 1999. Biomass yield of poplar after five 2-year coppice rotations. Forestry 72:157163. Prusinkiewicz, P.W. 1986. Graphical applications of L-systems. In Proceedings of Graphics Interface 86. Ed. W. Davis. Canadian Information Processing Society, Toronto, Ontario, pp 247253. Prusinkiewicz, P.W., W.R. Remphrey, G.C. Davidson and M.S. Hammel. 1994. Modeling the architecture of expanding Fraxinus pennsylvatica shoots using L-systems. Can. J. Bot. 72:701714. Rakocevic, M., H. Sinoquet, A. Christophe, C. Varlet-Grancher. 2000. Assessing the geometric structure of a white Clover (Trifolium repens L.) canopy using 3-D digitizing. Ann. Bot. 86: 519526. Remphrey, W.R., A.G. Bartlett and C.G. Davidson. 2002. Shoot morphology and fate of buds in relation to crown location in young Fraxinus pennsylvanica var. subintegerrima. Can. J. Bot. 80: 12741282. Ridge, C.R., T.M. Hinckley, R.F. Stettler and E. van Volkenburgh. 1986. Leaf growth characteristics of fast growing poplar hybrids Populus trichocarpa Populus deltoides. Tree Physiol. 1: 209216. Ripullone, F., G. Grassi, M. Lauteri and M. Borghetti. 2003. Photosynthesisnitrogen relationships: interpretation of different patterns between Pseudotsuga menziesii and Populus euramericana in a mini-stand experiment. Tree Physiol. 23:137144. Ross, J. 1981. The radiation regime and architecture of plant stands. W. Junk Publishers, The Hague, 391 p. Sinoquet, H. and X. Le Roux. 2000. Short term interactions between tree foliage and the aerial environment: an overview of modeling approaches available for tree structure-function models. Ann. For. Sci. 57:447496. Sinoquet, H. and P. Rivet. 1997. Measurement and visualization of the architecture of an adult tree based on dimensional digitizing device. Trees 11:265270. Sinoquet, H., S. Thanisawanyangkura, H. Mabrouk and P. Kasemsap. 1998. Characterization of the light environment in canopies using 3-D digitizing and image processing. Ann. Bot. 82:203212. Sinoquet, H., X. Le Roux, B. Adam, T. Amglio and F.A. Daudet. 2001. RATP, a model for simulating the spatial distribution of radiation absorption, transpiration and photosynthesis within canopies: application to an isolated tree crown. Plant Cell Environ. 24: 395406.
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CASELLA AND SINOQUET Tubby, I. and A. Armstrong. 2002. Establishment and management of short rotation coppice. Practice Note 7. Forestry Commission Research Information, Forestry Commission, Edinburgh, 12 p. Whitehead, D., J.C. Grace and M.J.S. Godfrey. 1990. Architectural distribution of foliage in individual Pinus radiata D. Don crowns and the effect of clumping on radiation interception. Tree Physiol. 7:135155. Zavitkovski, J. 1981. Structure and seasonal distribution of litterfall in young plantations of Populus Tristis #1. Plant Soil 60:409422.
Sonohat, G., H. Sinoquet, C. Varlet-Grancher, M. Rakocevic, A. Jacquet, J.C. Simon and B. Adam. 2002. Leaf dispersion and light partitioning in three-dimensionally digitized tall fescue-white clover mixtures. Plant Cell Environ. 25:529538. Stettler, R.F., H.D. Bradshaw, P.E. Heilman and T.M. Hinckley. 1996. Biology of Populus and its implications for management and conservation. NRC Research Press, Ottawa, Canada, 539 p. Takenaka, A. 1994. Effects of leaf blade narrowness and petiole length on the light capture efficiency of a shoot. Ecol. Res. 9: 109114.

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Tree Physiology 23, 11531170 2003 Heron PublishingVictoria, Canada

CASELLA AND SINOQUET

TREE PHYSIOLOGY VOLUME 23, 2003

P. deltoides P. nigra P. trichocarpa P. trichocarpa

TREE PHYSIOLOGY ONLINE at http://heronpublishing.com

CASELLA AND SINOQUET

TREE PHYSIOLOGY VOLUME 23, 2003

CANOPY ARCHITECTURE OF COPPICE POPLAR

TREE PHYSIOLOGY ONLINE at http://heronpublishing.com

CASELLA AND SINOQUET

TREE PHYSIOLOGY VOLUME 23, 2003

36 365 365 139 269

0.769 0.844 0.938 0.949 0.401

[10, 220] [11, 58]

nA1U1 a10 A1U1i a bS lA1U1 nM lI i lA1U1

d bS i nA1U1 d10A1U1 lA1U1 i nM

[0, 0.06] [0, 0.04] [16, 114]

[0, 1] [0, 1] [10, 220]

544 425 722 722 815

0.374 0.624 0.924

jSe1 jSenSe jSe1 jSenSe d10A1U1 2.152

33.00 90.00 16.00 90.00

0.671 0.543 0.148 0.171 0.010

4.983 4.507 1.43E3 1.78E3 5.27E5

16.24 15.89 1.89E7

21.31 22.01 3.140 1.628 1.641

36 312 312 110 476

0.000 0.821 0.978 0.965 0.453

[28, 305] [12, 50]

CANOPY ARCHITECTURE OF COPPICE POPLAR

nA1U1 a10 A1U1i a bS lA1U1 nM lI i lA1U1

d bS i nA1U1 d10A1U1 lA1U1 i nM

[0, 0.05] [0, 0.03] [6, 116]

650 465 747 737 652

0.550 0.686 0.964

jSe1 jSenSe jSe1 jSenSe d10A1U1

CASELLA AND SINOQUET

defined by six zenith divisions (i.e., six annuli with 15 intervals).

TREE PHYSIOLOGY VOLUME 23, 2003

CANOPY ARCHITECTURE OF COPPICE POPLAR

Clone Gho A1U2 11

A2U2 16 U2 15, 17 Clone Tri A1U2 11

TREE PHYSIOLOGY ONLINE at http://heronpublishing.com

CASELLA AND SINOQUET 0.990 0.997 0.080 0.696 0.502 0.090

92 1311 5676 559 276 105 386

84 423 4126 386 154 141

0.168 0.986 21.86

21.14 52.19 2.026 6.897

71.48 6.310 99.90 55.87

lA1U2 lA2U2 i/nA2U2 lL i/nM i/nM

[3, 48] [0, 35] [0, 1] [0, 7] [0, 1] [0, 1]

AL AL aL i / aL lP aPi jLi jLi

lA1U2 lA2U2 i/nA2U2 lL i/nM i/nM i/nM

TREE PHYSIOLOGY VOLUME 23, 2003

0.767 2.782 66.14 52 22 23.72

0.107 4.314 3.027 4.496 443.400

CANOPY ARCHITECTURE OF COPPICE POPLAR

TREE PHYSIOLOGY ONLINE at http://heronpublishing.com

CASELLA AND SINOQUET

A1U2 l nI DlI 2.4 aL

817 713 56 710 57 11 56 22 69 715 68 24 49 34 13 34 23 11 35

111 200 586 897 78 148 554 780

11932914 19314776 36157532 673815221 5502243 7292468 13813547 26598258