Tree Physiology 30, 1433–1447
doi:10.1093/treephys/tpq085
The effect of tree architecture on conduit diameter and frequency
from small distal roots to branch tips in Betula pendula, Picea abies
and Pinus sylvestris
ANNA LINTUNEN 1,3,† and TUOMO KALLIOKOSKI 1,2,†
1
Department of Forest Ecology, University of Helsinki, PO Box 27, FI-00014 Helsinki, Finland
2
Finnish Forest Research Institute, Vantaa Research Centre, PO Box 18, FI-01301 Vantaa, Finland
3
Corresponding author (anna.lintunen@helsinki.fi)
Received January 15, 2010; accepted August 29, 2010; handling Editor Ülo Niinemets
Summary We studied the effect of tree architecture on
xylem anatomy in three Betula pendula Roth., three Picea
abies (L.) H. Karst. and three Pinus sylvestris (L.) trees
(mean age 35 years). First, the analysis of conduit anatomy
in different tree parts showed that conduits tapered and their
frequency increased from roots (≥2 mm) to stem, from stem
to branches and further to leaf petioles in B. pendula.
Conduit anatomy in lateral and main roots, as well as lateral
and main branches, significantly differed from each other in
all the studied species. The increase in conduit diameter and
decrease in frequency from the pith to the bark were clear
aboveground, but variable patterns were observed below-
ground. In the leaf petioles of B. pendula, conduit diameter
increased and conduit frequency decreased with increasing
individual leaf area. Second, the results concerning the
scaling of conduit diameter were compared with the
predictions of the general vascular scaling model (WBE
model) and Murray’s law. The scaling parameter values at
the tree level corresponded with the predictions of the WBE
model in all the studied trees except for one tree of both
conifer species. However, the scaling parameter values
changed from one tree compartment to another rather than
remaining uniform inside a tree, as assumed by the WBE
model. The assumptions of the WBE model of a constant
conductivity ratio, constant tapering and an unchanged total
number of conduits were not fulfilled. When the conduc-
tivity ratio and relative tapering were plotted together, the
results aboveground corresponded quite well with Murray’s
law: the conductivity ratio increased when relative tapering
decreased. Our results support the theory that trees adjust
both their macro- and microstructure to maximize their
water transport efficiency, but also to prevent embolism and
ensure mechanical safety.
Keywords: conduit scaling, conduit tapering, tracheids,
vessels, xylem anatomy.
†
Both authors contributed equally to this work.
© The Author 2010. Published by Oxford University Press. All rights reserved.
For Permissions, please email: journals.permissions@oxfordjournals.org
Introduction
Xylem conduits are assumed to taper from the roots to the
base of the stem, and further to the leaves, in order to
achieve an optimal structure for vascular element function-
ing (e.g., Zimmermann 1978, 1983, Tyree and Zimmermann
2002). Concurrently, with decreasing cell diameter, cell
frequency was observed to increase in the same direction
(Zimmermann 1978, Tyree and Zimmermann 2002, De
Micco et al. 2008, Sellin et al. 2008). Sanio (1872) found in
his seminal work that tracheid diameter in Pinus sylvestris
increased from the stem apex downwards until the
maximum was reached, and then subsequently decreased
down to the stem base. Since Sanio, a basipetal increase
in conduit diameter has been reported by many other
researchers (reviewed by Tyree and Zimmermann 2002).
Zimmermann (1978, 1983) concluded that the reason for
conduit tapering was the control of water distribution. By
having the lowest conductivities in the minor branches at
the end of the flow path, the plant could control the distri-
bution of water regardless of the transport distance, thereby
resulting in similar values of total root–shoot conductance
despite different path lengths. In addition, tissue with
smaller conduits and a higher frequency in the distal parts
also minimizes the loss of conduction even in cases where
part of the tissue is deactivated because of drought, freezing
or other injuries (Tyree and Zimmermann 2002, De Micco
et al. 2008).
West et al. (1999b) stated in their model of the universal
scaling of vascular networks (later called the WBE model)
that plant vascular architecture conforms to the energy
minimization principle, leading to the tapering of conduits
from the stem base to the photosynthesizing compartments.
Conduit tapering was assumed to be continuous at the level
of the whole tree in order to minimize hydraulic resistance.
The model should, with only slight modifications, also be
able to account for the roots (West et al. 1999b). The WBE
1434 LINTUNEN AND KALLIOKOSKI
model employs several assumptions of tree architecture,
e.g., the crown is composed of a self-similar branching
pattern that is area preserving, the leaves are situated at the
terminal branching generation and the total conduit number
at each branching generation is constant. The branching
generation follows a developmental sequence starting from
the proximal segment (hereafter segment refers to a section
between two branching events) at the stem base as gener-
ation 1, its children as generation 2 and so on.
Murray’s law states that conduits should taper based on
the Hagen–Poiseuille value to maintain a constant relation-
ship between the volume flow rate and the sum of the
conduit radius cubed to maximize hydraulic conductance
with a given investment (Murray 1926, McCulloh et al.
2003, McCulloh and Sperry 2005). Assumptions of
Murray’s law lead to a hydraulic architecture where the ratio
of the conduit radius equals the conductivity ratio of adja-
cent segments (McCulloh and Sperry 2005). Murray’s law
does not assume the number of conduits to be fixed at each
level of branching generation, as the WBE model does, and
thus the maximum conductance also depends on how the
number of conduits changes between adjacent segments
(McCulloh et al. 2004).
Empirical results from previous studies seem to both
support and challenge the scaling theories. Murray’s law
has been found to apply to those tree parts that have no role
in mechanical support, whereas xylem conduits in support-
ing compartments seem to deviate from the law, the greatest
deviation being in coniferous trees (McCulloh et al. 2003,
2004, McCulloh and Sperry 2005). WBE-like tapering,
more (Anfodillo et al. 2006) or less (Coomes et al. 2007,
Mencuccini et al. 2007), has been observed in the stems of
both angio- and gymnosperms. However, a few recent
studies have accounted for the side branches (Atala and
Lusk 2008, Sellin et al. 2008) and root systems (Nygren and
Pallardy 2008) in addition to the main stem when testing
the WBE theory. When the roots and/or branches were
taken into account, the whole-tree conduit tapering pattern
was not continuous at the tree level but occurred rather as
leaps from one morphological level to the next (Atala and
Lusk 2008, Nygren and Pallardy 2008, Sellin et al. 2008). A
better congruence with the data was obtained and steeper
tapering in the roots compared with the stems was
suggested when the tapering ratio was allowed to change
along the pathway (Mencuccini et al. 2007). Conduit
tapering may vary between different species or growing
environments (Coomes et al. 2007).
Our first objective was to study the effects of tree archi-
tecture on conduit diameter and frequency in different
compartments of Betula pendula Roth., Picea abies (L.)
H. Karst. and P. sylvestris (L.) grown in early mature,
mixed boreal forest stands. Specifically, we determined the
effect of tree compartment, radial distance from the pith and
individual leaf area on conduit diameter and frequency. Our
second objective was to compare the theoretical predictions
of the WBE model and Murray’s law applied to our data
TREE PHYSIOLOGY VOLUME 30, 2010
set. As far as we know, our data set is the only one covering
the whole water pathway from distal roots to leaf petioles in
early mature trees.
Materials and methods
Study material and architectural measurements
The mixed study stands consisted of B. pendula, P. abies
and P. sylvestris and were located in Southern Finland
around the Hyytiälä Forestry Field Station (61°500 N, 24°
180 E, 160 m a.s.l.). Details of the field sites are presented in
Kalliokoski et al. (2008). The forest type was classified
according to Cajander’s (1949) system, in which stand ferti-
lity is assessed on the basis of the understorey vegetation.
The stands formed a site fertility gradient: the Oxalis–
Myrtillus site type (OMT) represented a fertile site, the
Myrtillus type (MT) a medium–high fertility site and the
Vaccinium type (VT) a medium–low fertility site. Three
individuals per species were sampled, i.e., the total number
of sample trees was nine. One B. pendula, two P. abies and
P. sylvestris were sampled at the VT site, one B. pendula at
the MT site, and one tree of each species was sampled at
the OMT site from the dominant trees (Table 1). The mean
age of the sample trees was 35 years (Table 2).
The study trees were felled and the crown of each tree
was cut into 1.5-m sections. The original north direction
was marked in each section for computationally reconstruct-
ing the tree architecture. In this study, tree architecture
refers to the spatial configuration of the system and was
described with branching frequency of the segments, length
of the segments, branching angles and topology, i.e.,
branching pattern (Fitter et al. 1991, Lynch 1995). Crown
architecture was determined from cut tree sections kept
standing in a wind shelter by digitizing, i.e., recording the
3D coordinates of five sample branches shoot by shoot.
Sample branches were systematically selected to represent
all the sections of the crown from the lowest branch to the
top. Digitizing was performed with the Polhemus Fastrak
system, equipped with a digitizing pen and Longranger
transmitter (Polhemus, Inc., Colchester, VT, USA). From
the digitized coordinates, we determined the exact location
and size of each digitized shoot, branch or stem part inside
the crown.
All the roots attached to the stump were exposed up to
the first branching event and one to two sample roots were
selected per tree. All segments of the sample roots were
excavated down to the preset minimum diameter of 2 mm.
Excavation was carried out by a pneumatic Soil-Pick tool
(MBW SP125; M-B-W Ltd, Bolton, UK). Details of the
root data are described by Kalliokoski et al. (2008).
We classified the tree segments according to the
Gravelius order (GO, MacDonald 1983) and distance from
the stem or branch apex. The GO assigns the same order
number to segments belonging to the same axis, and one
 XYLEM ANATOMY IN THREE BOREAL TREE SPECIES 1435
Table 1. Selected characteristics of mixed boreal stands situated in Southern Finland.
Forest site type Species Regeneration type Number of trees per hectare Basal area (m2 ha−1) Mean of stand

Height (m) D1.3 (cm)

OMT B. pendula Natural 530 6.2 14.0 12.6

OMT P. abies Planted 3150 6.5 5.8 6.0
OMT P. sylvestris Natural 270 4.7 14.4 14.4
OMT Others Natural 430 1.9 9.8 8.1
Total 4380 19.0 8.8 8.4
MT B. pendula Natural 480 5.0 12.9 11.5
MT P. abies Natural 3450 3.0 4.3 4.1
MT P. sylvestris Planted 830 10.7 11.5 12.2
MT Others Natural 930 0.2 3.0 1.3
Total 5690 19.0 8.2 8.0
VT B. pendula Natural 480 1.6 8.8 7.9
VT P. abies Natural 920 4.8 7.9 8.7
VT P. sylvestris Natural 390 3.3 8.7 10.4
VT Others Natural 110 0.2 5.9 5.7
Total 1900 10.0 8.2 8.8

D1.3, diameter at breast height, i.e., at 1.3 m from the ground surface.

Table 2. Aboveground characteristics of nine sampled trees

growing in mixed boreal forest stands.
Forest site Species Sample tree characteristics
type
Tree no. Age (years) D0 (cm) Height (m)

OMT B. pendula 6 40 24.0 18.7

OMT P. abies 4 33 16.7 14.0
OMT P. sylvestris 10 36 21.9 15.0
MT B. pendula 10 27 14.1 12.8
VT B. pendula 4 31 16.7 11.7
VT P. abies 2 37 13.3 7.7
8 45 18.0 10.8
VT P. sylvestris 3 32 19.2 10.3
7 33 19.1 10.8

D0, diameter of the stump; height was determined from the highest
point of the root collar or, when the root collar was completely
belowground, from ground level.

order number higher to lateral segments. The stem is the
main axis of order 1, order 2 is assigned to main branches
attached to stems, and so on. In belowground parts, the
main axis of the root is order 2, i.e., corresponding to the
main axis of aboveground branches, and lateral root
branches are one order higher. For the root and stem
samples, the apex of the leader shoot in the stem top was
considered a starting point for the distance calculations.
Correspondingly, for each branch sample, the starting point
was the axis apex.
Sampling
Vascular samples were taken to represent the whole water
transport pathway, fine roots and leaves excluded (Figure 1),
i.e., from the 2-mm lower diameter limit in the roots to the
Figure 1. Schematic figure of the applied sampling procedure. The
samples and sampling locations from different GOs are marked
with corresponding number of lines, i.e., GO 1 with one line, GO
2 with two lines and so on. We separated shallow and sinker roots
in sampling: sinker roots were denoted as root branches, the main
axes of which had a vertical angle of >45°.
final segment in the branches (leaf petioles in B. pendula,
distal shoots in conifers). Six stem discs were taken: the
first just above the root collar and one below each sample
branch (Figure 1). Analogously, five to six discs were taken
from the sample roots along the main axis (Figure 1). Three
discs were taken from the sample branches on each existing
GO in both the above- and belowground compartments
(Figure 1). Three leaves for leaf area measurements were
collected from each GO in each sample branch. The

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1436 LINTUNEN AND KALLIOKOSKI
Table 3. Sampling and imaging procedures in different tree compartments.
Compartment Sample Image

Root <1 cm Whole cross-section Continuous series of images, from pith to bark
Root >1 cm Three 1 cm2 pieces with even distances from pith to barka,b Images with even distances, from edge to edge of each pieces
Stem From second innermost ring, second outermost ring, and Three pictures, from each piece at even distances from early to
one in the middle of themb,c late wood
Branch Whole cross-sectionb Two pictures, one from early and one from late wood from
three growth rings selected at even distances from pith to bark
Leaf petiole Whole cross-section Whole cross-section
a
We did not measure specific growth rings in roots, because, in many cases, they were discontinuous and thus impossible to follow (cf.
Coutts 1986).
b
If reaction wood (compression wood or tension wood) was present in the sample, its effect was avoided by taking images from the normal
wood (Timell 1986).
c
This method enabled us to measure all the directions of conduit variation occurring in the stem: (i) radially from the pith to the bark,
(ii) vertically within a growth ring of a defined year and (iii) along the stem inside a growth ring of defined count from the pith (Burdon
et al. 2004).

measured petioles were connected to the collected leaves.
Leaves were photocopied and scanned, after which the area
was measured with Image-Pro Plus imaging software
(Media Cybernetics, Inc., Bethesda, MD, USA).
The woody tissue samples were stored in a freezer and
the petioles in liquid nitrogen. The sampling and imaging
procedures in different tree compartments are presented in
Table 3. Most of the woody samples were re-hydrated and
softened by boiling. Transverse sections of the aboveground
samples were taken with a sledge microtome (Reichert-Jung
Optiche Werke AG, Vienna, Austria) at a thickness of 100–
150 μm. The root samples were frozen and sectioned with a
cryomicrotome (Leitz 1516; Leica Microsystems, Wetzlar,
Germany), fitted with disposable microtome blades at a
thickness of 16 μm. The petioles were first embedded in
paraffin and then cut with the same microtome as the roots
at a thickness of 8 μm (Anderson and Bancroft 2002). All
the sections were stained with a mixture of safranin and
Alcian blue (ratio 1:2). The stained samples were then
rinsed with water, dehydrated in an ascending alcohol
series, rinsed with xylene and mounted in Canada balsam.
Xylem characteristics of the roots and petioles were
measured from the images captured with a SPOT Insight
B/W digital camera (Diagnostic Instruments, Inc., Sterling
Heights, MI, USA) attached to an Olympus BH-2 micro-
scope and from the stem and branches with an Olympus
ALTRA 20 colour camera attached to an Olympus CX31
microscope (Olympus Optical Co., Tokyo, Japan).
Image analysis
The images were analysed with Image-Pro Plus imaging
software (Media Cybernetics, Inc., Bethesda, MD, USA).
From the images of the crown samples, the area of conduit
lumen was first measured and then the conduit diameter
was calculated assuming the conduits to be circular, while
in the roots the lumen diameter was measured as the mean
of several distances passing through the cell’s centroid.
Conduit frequency was determined by calculating the
number of conduits within a defined area in the image. The
number of measured conduits per image was 101–613 in
the leaf petioles, 8–71 in the branches, 25–273 in the stem
and 4–145 in the roots. The total number of conduits
measured in the study was >100,000.
To estimate the total conductive area, we first calculated the
total basal sapwood area in each tree compartment. In the
branches and roots, this was done by summing up the basal
cross-sectional areas of all main branches/roots. We then multi-
plied the obtained sapwood area by the image-based tree- and
compartment-specific mean relative conductive area calcu-
lations. In calculations of mean relative conductive area, each
image represented an area of the sample defined by the dis-
tance from the branch or root pith.
Statistical analysis of xylem anatomy
Statistical analyses were carried out with SAS Statistical
Software v. 9.1 (SAS Institute, Cary, NC, USA). As the
data were hierarchical, the differences between tree compart-
ments were determined by a linear mixed model with a
nested structure for each species
y ¼ m þ bR R þ bS S þ bB B þ bP P þ ci þ d jðiÞ
þ ekðijÞ þ fmðijkÞ ; ð1Þ
where y is the conduit diameter or frequency, µ is the
general mean, R, S, B and P are representative of the fixed
effect of a class variable tree compartment, i.e., root, stem,
branch and petiole, respectively, and βR − βP are the
fixed-effect coefficients. The correlation structure for
B. pendula, P. abies and P. sylvestris among trees, roots and
branches, cross-sections and images in the data was
accounted for through random effects c (i = 3, 3, 3), d
( j = 36, 21, 22), e (k = 297, 230, 212) and f (m = 891, 873,
882), respectively. The Tukey–Kramer paired multiple
comparisons method was used to compare the tree

TREE PHYSIOLOGY VOLUME 30, 2010

 XYLEM ANATOMY IN THREE BOREAL TREE SPECIES
compartments. The same model structure was used to test
the differences in conduit diameter and frequency between
shallow and sinker roots. Sinker roots were denoted as root
branches, the main axes of which had vertical angle >45°
(Figure 1).
The effect of the radial distance from pith to bark on
xylem anatomy in different tree compartments, the differ-
ence in xylem anatomy between measured growth rings in
the stem and the effect of individual leaf area on xylem
anatomy in the petioles of B. pendula were studied with the
following mixed effects model structure:
y ¼ m þ bi Xi þ ci þ d jðiÞ þ ekðijÞ þ fmðijkÞ ;
where y is the dependent variable, Xi is the independent
fixed variable and βi is the corresponding slope. The
random variables are described with Eq. (1). The random
effects are c, d, e and f in the case of roots and branches,
e and f in the case of stems and c, d and e in petioles. The
Tukey–Kramer paired multiple comparisons method was
used to compare the growth rings in stems. Conduit diam-
eter and frequency were logarithmically transformed in the
analyses when necessary in order to normalize these vari-
ables and to reduce the heteroscedasticity of the residuals.
Scaling of conduit diameter
According to the WBE model, the conduit radius (a) tapers
as
akþ1
¼ nb=2 ;
ak
where k is the branching generation of a segment, n is the
number of segments formed in a branching point and b is a
scaling parameter. The theoretical value for the scaling par-
ameter b is 1/6 (West et al. 1999a). As it is impossible to
distinguish all the segments in a tree, especially in the
woody parts without visible forkings (due to shed
branches), we used the distance from the apex, l, instead of
k (Anfodillo et al. 2006). From the WBE theory, assuming a
volume filling network, it can be derived that the conduit
radius scales with distance from the apex as follows
(Anfodillo et al. 2006):
ak / lb=a ;
where α can be derived from the relationship between dis-
tance and tree diameter, D:
l ¼ Da
and should have a value of 2/3 based on the elastic simi-
larity model (Greenhill 1881, McMahon 1973, West et al.
1999a). As theoretical values of b and α are 1/6 and 2/3,
respectively, the theoretical scaling exponent b/α in Eq. (4)
is 1/4. Owing to the approximation of tree height in the
TREE PHYSIOLOGY ONLINE at http://www.treephys.oxfordjournals.org
1437
WBE model, an empirical correction factor of 0.84 is
needed when tapering of the conduit diameter is predicted
according to distance from the apex instead of segment
number (Anfodillo et al. 2006).
ak / l0:84b=a : ð6Þ
The exponent in Eq. (6) can be used to compare empirical
results with the WBE exponent (Anfodillo et al. 2006). We
estimated the exponent of Eq. (6) with the linearized model,
separately for each tree
ð2Þ y ¼ m þ bR R þ bS S þ bB B þ ekðijÞ þ fmðijkÞ ; ð7Þ
where y is the conduit diameter, R, S and B represent the
fixed effects of the compartment-specific logarithmic distance
from the apex, βR − βB are fixed-effect coefficients, i.e., the
exponent in Eq. (6), random effects being sample (e) and
image (f). We also studied how the conduit diameter changed
with distance from the apex in different growth rings of the
stem. This was analysed using the mixed model structure pre-
sented in Eq. (2), with random effects as in Eq. (7).
Murray’s law states that the volume flow rate, Q, and
conduit radius are related as (Murray 1926, McCulloh and
Sperry 2005):
X
Q¼ a3 : ð8Þ
The hydraulic architecture of a tree that conforms to Eq. (8)
leads to a xylem anatomy where the conductivity ratio of
adjacent segments, Kdist/Kprox, equals the relative conduit
ð3Þ tapering, T, between these segments
Kdist
¼ T: ð9Þ
Kprox
To examine whether our results corresponded to Murray’s
predictions in different species and tree compartments, we
calculated the potential conductivity K using the Hagen–
Poiseuille equation (Tyree and Zimmermann 2002) for each
segment in measured branching points. Then we compared
Kdist/Kprox, from distal roots towards the stem apex, in a
branching point with T. In the analyses, we used the diameter
and number of conduits obtained from the proximal end of
each segment to calculate the ratio of Kdist and Kprox and thus
accounted for the tapering taking place within a segment
ð4Þ (Soethe et al. 2007, Kalliokoski et al. 2010).
Results
ð5Þ
Xylem anatomy in different parts of a tree
Conduit diameter decreased and conduit frequency
increased from the roots to the stem and from the stem to
the branches in all three species, and further to the leaf
petioles in B. pendula (Figures 2 and 3). The compartment
1438 LINTUNEN AND KALLIOKOSKI

Figure 2. Microscopic images of xylem in different parts of a tree along the water transport pathway in (a) B. pendula, (b) P. abies and
(c) P. sylvestris. From right to left: distal root, root segment beside the stump, stem and side branches. In B. pendula, the rightmost picture is
a leaf petiole. The scale bars are presented in the figures. This figure appears in color in the online version of Tree Physiology.

level explained 83, 59 and 60% of the variation in conduit

diameter in B. pendula, P. abies and P. sylvestris, respect-
ively. Respective values for conduit frequency were 95, 81
and 85%. In the roots of B. pendula, the conduits were
smaller and more numerous in the lateral roots (GO > 2,
Figure 3) than in the main roots (GO = 2), but larger and
less frequent than in the stem. In conifers, the mean conduit
diameter in the lateral roots did not differ from the stem, but
the frequency was lower (Figure 3). Aboveground, the main
branches (GO = 2) had conduits with a larger diameter than
those in the lateral branches (GO > 2) in B. pendula and
P. abies, whereas there was no difference in P. sylvestris.
The conduit frequency was also higher in the main branches
than in the laterals in all the studied species.
There was a tendency for the shallow roots to have larger
conduits than the sinker roots. However, due to the small
number of sinker roots in our data (only 63 samples were
classified as sinker roots), this difference was not significant
in any of the species (P-values were 0.0596, 0.0686 and
0.3127 for B. pendula, P. abies and P. sylvestris, respect-
ively). Conifers also had a lower conduit frequency in
shallow roots than in sinker roots (P < 0.001).
In the roots, B. pendula and P. abies had significantly
larger and less numerous conduits in the outer wood than in
the core wood (P < 0.0001), whereas P. sylvestris displayed
the opposite trend (P < 0.0001). However, the variation Figure 3. Least-square means of conduit diameter (a) and conduit
in root xylem anatomy pattern from the pith outwards frequency (b) in sapwood of different tree compartments of
was large and the proportion of explained variation was B. pendula, P. abies and P. sylvestris. The error bars indicate the
extremely low in each species. The values of coefficient of 95% CI. Only early wood was measured in stems and branches of
the conifers. Different letters indicate significant differences
determination ranged from 0.01 to 0.08 in conduit diameter. between tree compartments within each species (Tukey–Kramer test
The coefficient of determination of conduit frequency was at P < 0.05). Analyses were performed with a mixed model using
0.33 for B. pendula, whereas for P. abies and P. sylvestris log-transformed values for conduit diameter (Eq. (1)).

TREE PHYSIOLOGY VOLUME 30, 2010

 XYLEM ANATOMY IN THREE BOREAL TREE SPECIES
Figure 4. Cross-sectional image of a 2.5-mm wide B. pendula root
showing the abrupt change in conduit size from the pith to the
bark. The scale bar is presented in the figure.
the values were 0.04 and 0.01, respectively. In many root
samples, the conduits were larger and less numerous in
some growth rings around protoxylem, but the size then
abruptly diminished and subsequently remained constant
(Figure 4). The proportion of latewood was also clearly
smaller than that observed in the aboveground parts. In
conifer sinker roots, the effect of radial distance on xylem
characteristics was opposite to that observed in the shallow
roots, i.e., the conduits were smaller (P < 0.0001) and more
numerous (P < 0.0001) in the outer wood than in the inner
wood in P. abies, and vice versa in P. sylvestris (P < 0.0001
for both conduit diameter and frequency). Conduit
diameter increased and frequency decreased from the pith
outwards in stems and branches of all the studied species
(P < 0.0001). In a stem disc of 10 cm diameter, the increase
in conduit diameter ranged from 61 to 102% and the
decrease in frequency ranged from 67 to 83%, depending
on the species. The conduit diameter and frequency differed
significantly (P < 0.05) between the second innermost
growth ring and the two outer rings, but were statistically
the same in the second outer ring and in the middle ring. In
1-cm diameter branches, the range of increase in conduit
diameter from the pith to the bark was from 45 to 97%, and
that in conduit frequency was from 52 to 96%, depending
on the species.
The estimated mean vessel diameter increased by 0.8 μm
when the individual leaf area increased by 5 cm2 (P <
0.0001, R 2 = 0.10) and the frequency decreased at the same
time by 997 vessels per square millimetre (P < 0.0001, R 2 =
0.55). From the smallest (150 mm2) to the largest measured
leaf (2120 mm2), the increase in the mean diameter of the
petiole vessels was
50% (Figure 5).
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1439
Figure 5. Mean conduit diameter in petioles versus leaf blade area
in B. pendula. Individual trees are denoted by different symbols.
The relationship was analysed with a mixed model (Eq. (2)).
Figure 6. The estimated total conductive area in different tree com-
partments of the studied species.
At the whole-tree level, the decrease in mean conduit
diameter from the most distal roots to the most distal
branches ranged from 40 to 80% (Figure 2), the decrease
being greatest in B. pendula. Conduit tapering was closely
accompanied by an increasing conduit frequency. At the
whole-tree level, our results suggested an acropetally
decreasing total conductive area (Figure 6). The total basal
conducting area values were the highest in roots (P < 0.1 for
B. pendula and P < 0.01 for the conifers). The mean values
of total conductive area seemed to decrease also from stem
to branches, but these differences were not significant in
any of the species due to the small number of sample trees
(Figure 6).
Scaling of conduit diameter
Conduit diameter increased with distance from the apex at
the whole-tree level (P < 0.05; Table 4, Figure 7). The
scaling exponents at the tree level were higher than pre-
dicted by the WBE model in all trees except for one
P. abies and one P. sylvestris (Table 4). Betula pendula had
higher scaling exponents than the conifers (Table 4), and
thus the predictions of the WBE model were better met in
B. pendula. However, scaling of the conduit diameter was
compartment specific rather than continuous, as suggested
1440 LINTUNEN AND KALLIOKOSKI
Table 4. Regression coefficients and 95% CIs for the model ln conduit diameter vs. ln distance from the apex (Eq. (7)).
Tree Species and N Intercept and 95% CI for the Scaling R2
compartment slopes slope exponent b
OMT 6 B. pendula 4412 2.8129*
Branches 0.7137* 0.4961–0.9313 0.600
Stem 0.3713* 0.3308–0.4118 0.312
Roots 0.8164 0.1876–1.4453 0.686
Tree level 2.9107* 0.82
0.3578* 0.3280–0.3877 0.301
MT 10 B. pendula 4568 2.5721*
Branches 0.5542 0.3564–0.7522 0.466
Stem 0.5935* 0.5537–0.6334 0.499
Roots 0.3342 −0.2930–0.9615 0.281
Tree level 2.5664* 0.96
0.5848* 0.5603–0.6092 0.491
VT 4 B. pendula 4046 2.8982*
Branches 0.6067 0.3919–0.8216 0.510
Stem 0.4830* 0.4362–0.5299 0.406
Roots 0.1257 −0.2858–0.5373 0.106
Tree level 2.9449* 0.91
0.4365* 0.4100–0.4631 0.367
OMT 4 P. abies 8254 2.4252*
Branches 0.5804* 0.3155–0.8453 0.488
Stem 0.2945* 0.2413–0.3477 0.247
Roots 0.7589 −0.0054–1.5233 0.637
Tree level 2.4860* 0.81
0.2985* 0.2660–0.3311 0.251
VT 2 P. abies 5895 2.4653*
Branches 0.6230 0.2499–0.9962 0.523
Stem 0.3302* 0.2712–0.3893 0.277
Roots 0.4304 0.1108–0.7501 0.362
Tree level 2.5103* 0.85
0.3275* 0.2955–0.3595 0.275
VT 8 P. abies 10254 2.4549*
Branches 0.3815 0.0143–0.7488 0.320
Stem 0.2283* 0.1697–0.2869 0.192
Roots −0.3936 −1.1366–0.3493 −0.331
Tree level 2.5008* 0.52
0.1781* 0.1437–0.2125 0.150
OMT 10 P. sylvestris 4579 2.6940*
Branches 0.4410 0.1789–0.7032 0.370
Stem 0.2707* 0.2239–0.3176 0.227
Roots 0.7579 0.1168–1.399 0.637
Tree level 2.7220* 0.81
0.2873* 0.2579–0.3167 0.241
VT 3 P. sylvestris 3402 2.5799*
Branches 0.6044* 0.2490–0.9599 0.508
Stem 0.3161* 0.2455–0.3866 0.266
Roots 0.5833 −0.4050–1.5717 0.490
Tree level 2.6369* 0.70
0.3091* 0.2610–0.3571 0.260
VT 7 P. sylvestris 12148 2.6379*
Branches 0.3834 0.0527–0.7142 0.322
Stem 0.1943* 0.1425–0.2462 0.163
Roots 1.3115* 1.0772–1.5458 1.102
Tree level 2.5496* 0.55
0.3074* 0.2609–0.3539 0.258
Values marked with an asterisk are significant with P < 0.05. The scaling exponent is calculated by multiplying the slope by 0.84 (Anfodillo
et al. 2006). The tree level values of coefficients of determination (R 2) describe the goodness of fit between the predictions of the WBE
model and the observations.

TREE PHYSIOLOGY VOLUME 30, 2010

 XYLEM ANATOMY IN THREE BOREAL TREE SPECIES
Figure 7. Mean conduit diameter in sample trees from the VT site
plotted against distance from the apex; (a) B. pendula, (b) P. abies
and (c) P. sylvestris. Roots, stems and branches are denoted with
different symbols. Tree-level WBE theory lines are drawn based on
a theoretical exponent of 1/4 and intercept values are taken from
the data in Table 4.
by the WBE model (Table 4, Figure 7). The compartments
differed in the slope values statistically (95% confidence
intervals (CI) did not overlap) in only two out of the nine
trees, but in most cases the likelihood of different compart-
ments having similar slope values was really small (the
95% CIs of the slope values overlapped only slightly).
Scaling exponents were generally higher in roots and
branches than in stems, and four out of the five highest
slope values were observed in roots (Table 4).
In the second outer growth ring, the conduit diameter
also increased with increasing distance from the apex in the
stem of each B. pendula tree, in one P. abies and in one
P. sylvestris (P < 0.05; Figure 8). Conduit diameter in the
second inner growth ring of stem, in turn, decreased with
increasing distance from the apex in four out of six conifer
trees (P < 0.05), but did not change in B. pendula trees
(Figure 8). In the middle ring, there was no significant
TREE PHYSIOLOGY ONLINE at http://www.treephys.oxfordjournals.org
1441
Figure 8. The effect of distance from the stem apex on mean
conduit diameter in different growth rings in sample trees growing
on the OMT site.
change in conduit diameter in five out of the six conifer
trees, whereas in two out of the three B. pendula there was
an increase in the vessel diameter with distance from the
apex (P < 0.05; Figure 8).
When the conductivity ratio Kdist/Kprox and relative taper-
ing T in a branching point were plotted together with
Murray’s law and the WBE optimum, the aboveground
results corresponded rather well to Murray’s law, especially
in the stem (Figure 9). In branches, the conductivity ratio
was a little below the one suggested by Murray’s law with a
given tapering ratio (Figure 9). In roots of each species,
however, the observed conductivity ratio was much higher
than that predicted by Murray’s law. The furcation number,
a ratio of the total number of conduits in distal and proximal
segments, was >1.00 throughout the trees, except in the
roots of B. pendula (average 1.00) and stems of P. abies
(average 1.04). Overall, in roots (1.00–1.17) and stem
(1.04–1.18), the values were closer to 1.00 than in branches
(1.24–1.29, Figure 9). Stems were closest to the WBE
tapering optimum, as conduits in branches tapered more
1442 LINTUNEN AND KALLIOKOSKI

Figure 9. Conductivity ratio (Kdist/Kprox) plotted against relative tapering in different species and tree compartments. Ratios were taken
between distal and proximal segments (towards the apex) in a branching point, including the change inside the segment, i.e., observations
from the beginning of the distal and proximal segments were used. Aboveground, conduit diameter (R 2 = 0.86–0.95) and frequency (R 2 =
0.98–1.00) were estimated from the segment diameter separately for each tree. Belowground, relative conduit tapering and conductivity were
only calculated for those branching points in which all the segments were measured. Murray’s law holds when the conductivity ratio equals
the relative tapering (solid line). In the figure, the black circle represents the value obtained with the assumptions of the WBE model, i.e.,
two child segments, a tapering exponent of 1/6 and constant conduit frequency.

strongly, whereas the conductivity ratio was closer to that
predicted by the WBE in branches (Figure 9).
Discussion
Xylem anatomy in different parts of a tree
We studied xylem anatomy in different tree compartments
of B. pendula, P. abies and P. sylvestris in order to assess
the effect of tree architecture on xylem characteristics.
Conduits were largest in the roots and tapered to the stem
and from there on, to the branches in all species, and further
to the leaf petioles in B. pendula. Concurrently, conduit fre-
quency increased from the roots to the branches. The tree
compartment as an explanatory variable explained 59–83%
of the variation in conduit diameter and 81–95% of that in
conduit frequency, depending on the species. The maximal
observed decrease in mean conduit diameter from the roots
to the branches ranged from 67 to 87% and the frequency
increased from 58 to 95%, depending on the tree. The
decrease of conduit diameter was greatest in B. pendula.
These results correspond to earlier findings of an acropetally
decreasing cell diameter between successive tree compart-
ments (Fegel 1941, Bailey 1958, Bhat 1981, Baas 1982,
Zimmermann 1983, Oliveras et al. 2003). Owing to the
assumption made of conduit circularity aboveground, our
results include a slight overestimation of conduit diameter
in stem and branches. Thus, the difference in conduit diam-
eter between above- and belowground parts may be even
larger than the results suggest. Summing up the two
phenomena, i.e., the decrease in conduit diameter and the
increase in frequency, the total water conductive area
decreased from roots to stem and from stem to branches.
One explanation could be that top-heavy structures would
be mechanically unstable (McCulloh and Sperry 2005).
However, the approach we used in calculating the total
basal conducting area for each tree compartment is only
valid if the pipe model assumption of area-preserving
branching holds true (Shinozaki et al. 1964). This seems to
be questionable, at least in the roots (Kalliokoski et al.
2010). We did not also account for the effect of heartwood
in stem. In roots and branches, no heartwood was observed.
The observed difference between root types, i.e., larger
and fewer conduits in shallow roots than in sinker roots, has
also been observed in other studies (Bannan 1941, Fahn
1964, Patel 1965, Fayle 1968, Zimmermann 1983, Sieber
1985, Petit et al. 2009). Under humid conditions, the most
important role of sinker roots may be to anchor the tree
more firmly in the soil (Reubens et al. 2007). Thus, small
conduits in sinker roots may be a response to mechanical
demands. Differences were also observed in conduit dia-
meter and frequency between branch hierarchies. Conduit
diameter was larger in the main branches of B. pendula and
P. abies than in the lateral branches, but there was no stat-
istically significant difference in P. sylvestris. Conduit fre-
quency was also lower in the lateral branches of the
conifers, which led to a decreasing total conductivity from
the main to the lateral branches per given branch
cross-sectional area. A reason for the decreasing conduit
frequency could be, for example, an increase in cell wall
thickness from the main to the lateral branches or change in
the relative proportion of xylem rays. Our results suggest
slight top-heaviness in the sapwood area of branches (data
not shown), which implies a complimentary mechanism for
sustaining total conductivity. Conduit diameter has been
reported to decrease towards higher branching hierarchies in
other studies on B. pendula (Atala and Lusk 2008, Sellin
et al. 2008) and in Populus deltoides Bartr. ex Marsh.

TREE PHYSIOLOGY VOLUME 30, 2010

 XYLEM ANATOMY IN THREE BOREAL TREE SPECIES
(Nygren and Pallardy 2008). Conduit frequency was found
earlier to increase in B. pendula from the main branches to
lateral branches (Sellin et al. 2008), whereas there was no
statistical change in our study. Total conductivity decreased
laterally in our study and also in the study of Sellin et al.
(2008). A functional explanation for the differences in
conduit anatomy between the main and lateral branches
could be apical control, i.e., the inhibition of lateral branch
growth by shoots above it (Wilson 2000). Rather than a
bushy crown architecture, apical control leads to a dominant
main stem axis that is characteristic of the species studied,
and to the prioritization of the stem and main branches, e.g.,
in the case of drought, the lower water potential in the
branches ensures the continuation of water flow in the stem
(Wilson 2000). Our results suggest that the same mechanism
between main and lateral axes of the branches may exist.
The apparent similarity between xylem anatomy and apical
control is probably a result of apical control due to growth
inhibition in the lateral branches rather than a cause of it,
since it has been shown that an increase in xylem cross-
sectional area follows, but lags far behind, release from
apical control (Spicer and Gartner 1998, Wilson 2000).
Aboveground, conduits increased in diameter and
decreased in frequency from the core to the outer wood.
Our observation agrees with the results of Saranpää et al.
(2000). The phenomenon is usually related to the effect of
cambium ageing (Gartner 1995, Spicer and Gartner 2001).
However, there was large variation in the root data and, in
the roots of P. sylvestris, conduit diameter even decreased
from the core to the outer wood. Variable xylem patterns in
a radial direction have also been reported in earlier root
studies (Bannan 1941, Peterson et al. 2007, Christensen-
Dalsgaar et al. 2008). Consistent with our observation con-
cerning P. sylvestris, Bannan (1941) found that most of the
sapwood area in the core wood of distal parts of lateral
roots in mature trees was early wood consisting of the
largest tracheids. Accordingly, vessel size decreased from
the core to the outer wood in the distal part of buttress roots
of three tropical tree species (Christensen-Dalsgaard et al.
2008): Xylopia nitida (Dunal), Tachigali melinonii (Harms)
Zarucchi & Herend. and Pradosia cochlearia (Lecomte)
T.D.Penn. We also observed this radial decrease in some
of the B. pendula and P. abies samples (Figure 4). The
decrease in conduit diameter could be interpreted as a result
of mechanical loading transmitted over a greater distance
from the stem base as the size of the tree increases, i.e., a
‘non-supporting root’ with large conduits has become a
‘supporting root’ with small conduits (Fayle 1968). Thus,
the conclusion that the radial anatomical patterns are not a
passive function of cambial ageing, but rather a response to
local mechanical loading (Christensen-Dalsgaard et al.
2008), is supported by our data. The maturing vascular
cambium is not constrained to produce only one radial
pattern of wood properties (Peterson et al. 2007).
Conduit diameter in the leaf petioles of B. pendula
increased and conduit frequency decreased with increasing
TREE PHYSIOLOGY ONLINE at http://www.treephys.oxfordjournals.org
1443
individual leaf area. This is a logical outcome, since a
larger leaf needs larger vessels in order to respond to the
higher transport demands created by higher quantum flux
absorbance and evaporation. At tree level, conduits are
smallest and most numerous in the leaves and petioles. The
majority, up to 80%, of the total resistance of the water
pathway has been proposed to be the result of non-vascular
transport and transport in small conduits in fine roots and
leaves (Tyree and Zimmermann 2002). The total water con-
ducting area of tree petioles was slightly higher than in the
branches (Figure 6). Petioles are mechanically supported by
turgor pressure and collenchyma, and conduits are thus
not required to function as mechanical support (McCulloh
et al. 2003). A high total conducting area in the petioles
(Figure 6) does not necessarily mean a top-heavy crown
structure, as the density of the petiole biomass is far less
than that of branch wood.
Scaling of conduit diameter
Mean conduit diameter increased with increasing distance
from the apex in all three species, and the values of scaling
exponents were higher than the proposed theoretical value
of 1/6 corresponding to the WBE model (West et al.
1999a). However, the scaling of conduit diameter was
found to be compartment specific rather than uniform at the
whole-tree level. Scaling of conduits was evident both in
roots and branches, although there was large variation
between trees in the roots. Our results are consistent with
other studies in which decreasing conductivity has been
found along the branch length (Sanio according to Bailey
and Shepard 1915, Tyree and Zimmermann 2002, Sellin
et al. 2008). In the stems of the conifers, the minimum
value of the scaling exponent proposed by the WBE model
was exceeded in four trees out of the six. The overall
pattern of the conduit mean diameter in the stem as a func-
tion of height resembled that found by Sanio in P. sylvestris
(Bailey and Shepard 1915): the mean conduit diameter of
the whole cross-section in most cases increased from the
stem apex downwards, reached the maximum at an inter-
mediate height and then decreased. Recently, tapering in the
stem of B. pendula has been reported to be vertically negli-
gible (Sellin et al. 2008), or conduits have been reported to
taper only in the upper half of the crown (Westhoff et al.
2009). However, as we observed large conduits along the
whole stem, their effect on the total transport resistance
could be negligible compared with that of the distal parts.
This makes it possible for the branches at various heights
within the crown to transport relatively similar amounts of
water, regardless of the transport distance (Tyree and
Zimmermann 2002). This observation also lends support to
the conclusion of Sellin et al. (2008) that trees may adjust
the ratio of the sapwood to leaf area in response to increas-
ing transport distance rather than change the xylem anatomy
in the stem.
1444 LINTUNEN AND KALLIOKOSKI
Acropetal tapering of conduits may lead to the energy
minimization principle; trees constructed in this way would
maximize the hydraulic conductance of the xylem with
minimal investment in vascular tissue (West et al. 2000).
This interpretation is, of course, a simplification of complex
phenomena, because the anatomical changes in the branch
junctions (Schulte and Brooks 2003) and resistance in the
radial direction (Comstock and Sperry 2000) have not been
accounted for. Conduit anatomy is a trade-off between
hydraulic efficiency, avoidance of cavitation and mechanical
demands (Tyree and Zimmermann 2002, Sellin et al. 2008).
The differences in conduit scaling between different com-
partments inside a tree may reflect mechanical demands and
the minimization of embolism and its consequences rather
than a response to the actual transport distance, i.e., the
tapering would be more a solution for the optimization of
performance while ensuring safe hydraulic architecture
(Zimmermann 1983, Atala and Lusk 2008).
Steeper anatomical scaling has been predicted for roots
than stems, because of the lower construction costs of the
thin-walled root conduits (Mencuccini et al. 2007). This
proposal was supported by our results as in six of the nine
trees the slope value in roots overlapped that in stem
(Table 4). Roots have lower requirements for mechanical
strength and a lower risk of implosion because of the lower
negative water potentials. Embolism may also be more
easily reversible in roots than in other parts of the tree
because of positive root pressures (Tyree and Sperry 1989),
and the area projected to the soil surface through which the
mechanical loading of a tree is transmitted is relatively
restricted compared with total root extension (Laitakari
1927, Stone and Kalisz 1991, Nicoll et al. 2006, Kalliokoski
et al. 2008).
The high production of reaction wood near the branch
base (Timell 1986) and the fact that the stem has to keep
the tree upright and resist bending caused by the pushing
forces of prevailing winds (Holbrook and Putz 1989, Kern
et al. 2005) indicate that the mechanical demands may
detract the scaling of conduit diameter further from the
optima of water transportation. However, as we excluded
reaction wood from our analysis aboveground, our results
on conduit scaling from the apex may reflect more the
effect of hydraulic demands. The effect of possible flexure
wood (e.g., Kern et al. 2005) was not taken into account in
our analysis, since unlike reaction wood, flexure wood is
not as clearly identifiable in a transverse section.
Regarding the differences between the three species,
B. pendula had the highest scaling coefficient at the whole-tree
level. As the sapwood of B. pendula consists of vessels and
fibres, its conduit diameter may respond to water transport
with fewer constraints because the mechanical demands are
directed to fibres. Conduit tapering has generally been
observed to be more pronounced in angio- than gymnosperms
(Mencuccini et al. 2007). On the other hand, the need to
decrease conduit diameter distally for hydraulic safety maybe
more pronounced in B. pendula than in the conifers, in which
TREE PHYSIOLOGY VOLUME 30, 2010
water is transported from one cell to another via bordered pits
containing a torus-margo structure. This structure efficiently
prevents embolism (Choat and Pittermann 2009) but, on the
other hand, may account for up to 50% of the total transport
resistance (Choat et al. 2008). The relationship between the
torus diameter and pit aperture diameter increases towards the
tree top, which means increasing safety against cavitation, but
decreased water conductance (Domec et al. 2008). The studied
models do not consider pit resistance, and it can thus be
questioned whether these models are suitable for conifers in
the first place.
An acropetal conduit tapering pattern in tree stems has
usually been observed in studies on a single growth ring
(Zimmermann 1978, Anfodillo et al. 2006, Coomes et al.
2007). The acropetal decrease in cell size and increase in
frequency have been linked to distance from the auxin
source (Aloni 1987, 2001); acropetally increasing auxin
levels lead to higher cell density and faster cell differen-
tiation rates, and thus to larger conduits in basipetal tree
parts due to slower cell differentiation and thus a longer
time available for cell expansion. We observed acropetal
tapering in conduit diameter in the second outermost
growth ring in each B. pendula and in one individual of
both conifers. In a second innermost growth ring with a
fixed cambium age, conduit diameter did not change acro-
petally in B. pendula, but increased in four out of the six
studied individual conifer trees. In diffuse-porous trees, 70–
90% of the sapwood area conducts water (Gebauer et al.
2008). Owing to the fourth power relationship between
conduit radius and conductance in the Hagen–Poiseuille
equation, the majority of the water volume is transported in
the outer growth rings where the largest conduits are located
(Wullschleger and King 2000, James et al. 2002,
Nadezhdina et al. 2002), whereas smaller conduits near the
pith primarily act as water storage sites (Goldstein et al.
1998). The latter are also safer pathways in the case of
drought or other injuries, because the largest conduits in the
outer growth rings are the most vulnerable to embolism
(Meinzer et al. 2001).
Murray’s law applied rather well to our observations in
the stems and branches, whereas in the roots, the conduc-
tivities were much higher than proposed at a given relative
tapering. Unlike the WBE model, Murray’s law allows the
furcation number of conduits to change inside a tree. The
average furcation numbers 1.00–1.29 found in this study are
in line with those reported earlier by McCulloh et al. (2004)
in non-supporting structures, which is quite a surprise,
because values >1.00 should only be observed in tree com-
partments free of any mechanical tasks. Furcation numbers
>1.00, together with an optimal relationship of Murray’s
law between relative tapering and the conductivity ratio,
lead to a top-heavy structure that is not mechanically stable
(McCulloh et al. 2004). In our data, the furcation number
was >1.00 especially in branches, but the conductivity
ratio was below Murray’s prediction at the given relative
conduit tapering. Nevertheless, the results suggest small
 XYLEM ANATOMY IN THREE BOREAL TREE SPECIES
top-heaviness in branches, but at the tree level between the
stem, branches and petioles the situation was the opposite.
Inside roots, conductivity increased from proximal to
distal segments, which is an interesting finding. A top-heavy
structure is not favourable in the aboveground woody parts
of a tree, whereas more biomass near the stem base and the
root collar is needed belowground in order to increase the
mechanical strength of the root system. Root values regard-
ing the Murray analysis include considerable uncertainty,
because our data set had only 37 such branching points
with all the proximal and distal segments measured.
Relative tapering values in the analysis of Murray’s law are
not comparable to the scaling exponent values in Table 4,
because here tapering was studied locally in each branching
point, whereas in the analysis of the WBE model the
scaling values were derived for each segment in relation to
the distance from the apex.
Both the WBE model and Murray’s law are universal
models and thus simplicity and assumptions are their
strengths. As the architecture of only a few tree species
fulfils the assumptions of the WBE model (McCulloh and
Sperry 2005), it is not unexpected to have results that differ
from its predictions if dealing with specific tree species.
Thus, part of the observed deviations from the WBE model
predictions in this study may be explained as a result of the
different crown structures from those assumed in the WBE;
the xylem anatomy was affected by the differences between
main axes and laterals, and the terminal units were not
located on a specific branching generation. Our results also
suggest that inclusion of the roots needs more than minor
modifications as suggested by West et al. (1999b).
Furthermore, Mäkelä and Valentine (2006) have shown that
the parameter b value of 1/6 does not minimize the hydro-
dynamic resistance. Murray’s law allows the furcation
number of conduits to change, which seems to be required
when moving from theory to studying real plants. However,
neither of these models accounts for any mechanical
demands, which should be kept in mind when interpreting
the observed results.
Conclusions
We found that conduit diameter decreased and frequency
increased from the roots to the stem, from the stem to the
main branches, from the main branches to the lateral
branches and further to the leaf petioles in B. pendula.
Conduits increased in diameter and decreased in frequency
aboveground from the pith to the bark, whereas below-
ground there was more variation in the radial pattern of
conduit anatomy. The scaling of conduit diameter also dif-
fered between compartments inside the trees. The assump-
tions of the WBE model of a constant conductivity ratio,
constant tapering and an unchanged total number of con-
duits were not fulfilled. Thus, our results are in line with
TREE PHYSIOLOGY ONLINE at http://www.treephys.oxfordjournals.org
1445
some earlier findings that while the WBE model has gained
considerable support in a general sense, the model could
not give a realistic description of the transport system of
specific species. Murray’s law was not tested as extensively
as the WBE model in this study, but it seemed to be flexible
enough to give tools for analysing and comparing different
compartments inside a tree.
Acknowledgements
We gratefully acknowledge the help of our supervisors Risto
Sievänen, Pekka Nygren and Pekka Kaitaniemi in data analysis and
writing the manuscript. We thank Pekka Saranpää for giving us the
possibility to use the wood anatomy laboratory of Metla. We also
thank Claire Baudelot and Camille Piquet for their skilled labora-
tory assistance, Teemu Hölttä for constructive comments on the
manuscript and appreciate the help received from Tuula Jyske,
Riikka Piispanen and Irmeli Luovula in different stages of the
laboratory work. Thank you also for the anonymous referees for
the constructive comments of the manuscript.
Funding
The study was funded by the Academy of Finland (Project
210875) and by the Finnish Graduate School in Forest Sciences.
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