Leonardo Was Wise

Trees Conserve Cross-Sectional Area Despite Vessel Structure

Rizwan Aratsu

Abstract

Beginning with Leonardo da Vinci's assertion that trees conserve total cross-sectional area
across every branching point, I tested ten species of trees in the vicinity of Princeton, New
Jersey, to see if they do indeed adhere to the rule of conservation as asserted by the Italian
master and those who followed him. Based on my review of the literature, I expected to find
that trees would either conserve area or not depending on the porosity of their wood to
water. To my surprise, I found that all ten species conserve cross-sectional area in
approximately the same way despite large differences in porosity. In particular, their twigs
roughly doubled in cross-sectional area across each branching while their larger branches
approximated area-preservation, as Leonardo had predicted. Rather than precisely
preserving area, the trees actually tended to increase in area ever so slightly as I moved
from trunk to twig tips. For this reason, I describe a conical model of tree form originated by
Horn (1998, in press), which may estimate the volume of a tree more accurately than the
traditional cylindrical model.

Introduction
Leonardo's Place in History

Figure 1. In his notebook, Leonardo da Vinci made this sketch depicting the branching pattern of trees. He depicted that
the total thickness of branches along each of the arcs would e qual the thickness of the trunk. (Richter 1939, plate 27)

Many observers of nature -some scientists, some poets, some both- have attempted to
explain the complex structure of trees. One of the most perceptive of these descriptions was
made in the 15th Century by Leonardo da Vinci who observed that"all the branches of a tree
at every stage of its height when put together are equal in thickness to the trunk" (Richter
1939, see Figure 1). In essence he was saying that the structure of trees is simple;that the
cross-sectional area of a tree stays the same from its base to the tip of its smallest branch.
It was such a beautifully articulated observation that it became, for lovers of trees, the basis
for our understanding of branching structure until today.
The Pipe Model Theory and its Critics
Leonardo's theory has not yet been explicitly tested. It was simply assumed to be true,
redefined by modern scientists, and then built upon. Shinozaki et al (1964a & b)
reformulated this theory by conceptualizing a tree as a bundle of unit pipes, each of which
supplies a particular amount of leaf area with water. They found a direct correlation
between the total cross-sectional area at any horizontal level and the leaf mass above that
level. From that time on, the analysis of branching structure has, for the most part, shifted
away from structural allometry (scaled physical measurements) towards vascular allometry
based on this "Pipe Model Theory" of Shinozaki et al.
The Pipe Model has significant implications for conservation biology. One of its potential
uses is for estimating the biomass of forests, a common practice among foresters interested
in lumber as well as scientists who want to know about the dynamics of carbon in the
ecosystem.

Figure 2. An illustration of a tree that is compressed into a cylinder so that we can estimate its volume.

Rather than trying to measure the volume of a tree in its natural form, we can conceptually
compress it into an upright cylinder (see Figure 2) if three key assumptions hold true. The
most basic assumption is, as the Pipe Model asserts, that the total cross-sectional area is
conserved across each branching. The second, which applies only to trees that branch
dichotomously (into two even branches), is that all the branches of a given diameter should
be of equal length. The final assumption, which applies only to excurrent branching patterns

(like a bottle-brush), is that branch length should increase proportionately as we move from
base to canopy. If these three assumptions hold true, then we can conceptualize the tree as
a cylinder whose base is the area of the cross-section of the trunk, and whose height is the
height of the tree (again see Figure 2). To calculate the area of the base of that cylinder we
need only measure the diameter at breast height (DBH). To find the height of the cylinder,
we measure the height of the tree using any of a variety of estimation methods. Both these
measurements are easy to make, and the shortcut provides an acceptable approximation of
the true volume of the tree (Robichaud & Methven 1992; Nygran et al 1993). Best of all,
this method greatly increases the efficiency with which biologists can estimate forest
biomass.
The Pipe Model is intuitively appealing once we understand the basics of water transport in
trees. Trees must draw tremendous amounts of water from their roots up to their leaves in
order to transport nutrients, dissipate heat, and maintain turgor pressure for support. The
driving force of this water pump is the transpiration that occurs in the leaves, which draws
up a continuous column of water through the xylem vessels (Zimmermann 1983). Since a
plant's competitive advantage depends in large part upon its success in fixing carbon (i.e.
the production of sugar and wood), and carbon fixation is limited by water accessibility, it is
reasonable to say that a plant's growth is limited by its ability to get water to its leaves. For
this reason, there should be a close correlation between a tree's conductive ability and its
leaf biomass or leaf number.
However, upon closer examination, the Pipe Model has proven inadequate to describe the
structure of real trees, though its essence still serves as the starting point for such studies.
In particular, criticisms have focused on two assumptions made by the model. Shinozaki et
al assumed that the xylem vessels of trees act like ideal pipes and that flow through them
follows Poisseuille's law (in Tyree & Ewers 1991) which, in essence, says that the flow
through a pipe is proportional to its radius, raised to the fourth power (i.e. flow ( radius4).
In other words, if we double the radius of a pipe, we increase the flow rate by a factor of
sixteen. While Poisseuille's law does help us to understand the gross differences in flow
between trees with different vessel sizes, it fails to describe the details of flow accurately for
four reasons. First, vessels are not perfectly cylindrical. Second, there is too much variation
in the size of vessels of a single tree to make such a sweeping generalization about all
vessels. Third, vessels do not maintain a constant diameter along their entire length; rather
they taper and expand in strategic places (Zimmerman 1978, 1983; Tyree & Ewers 1991;
Tyree & Alexander 1992). Finally, vessels are not actually continuous at all; rather they are
made up of shorter units that are juxtaposed end to end (see Figure 3).
Shinozaki et al's second assumption was that trees conduct water through all existing xylem
vessels regardless of the vessels' ages. Much research has been done to show that this is, in
fact, not true. When a tree no longer has any need for its old vessels, or when they become
too costly to maintain, it seals them with various resins and gums, forming what are called
tyloses (pronounced ty-LOW-sees) (Zimmermann 1983; Cochard & Tyree 1990; Tyree &
Ewers 1991). This phenomenon is familiar to anyone who has sawn through red cedar and
seen and smelled its resinous heartwood. In some species, the vessels are sealed, not
because the tree seals them physiologically, but because circumstances demand it. In these
trees, air comes out of solution when the water in the xylem freezes in the winter, leaving
deadly air bubbles called embolisms. Once such air bubbles form, the tree can no longer use
the embolized vessels unless it can reabsorb the bubbles. If not, it must make a new set of
pipes for the coming year (Tyree & Alexander 1993; Sperry et al 1994).

Figure 3.From left to right, non-porous, diffuse porous, and ring-porous vessels are illustrated here. (Zimmermann 1963,
p. 5)

Whether a tree forms tyloses or embolisms depends largely on its inherent vessel size (see
Figure 3). Trees with large vessels such as Americal elms (Ulmus americana) and
oaks (Quercus spp.) embolize annually as a regular part of their life cycle (Zimmerman
1983; Cochard & Tyree 1993). Each winter, the water in their vessels freezes, and the
dissolved air vaporizes, collecting in large bubbles. When the ice thaws in the spring, the
bubbles cannot fully dissolve because of their large size. If the leaves were to begin
transpiring, these bubbles would prove deadly, for they would expand and break the
continuity of the water column, thereby starving the leaves. To avoid this effective suicide,
these trees add a fresh layer of xylem early in the season, before the leaves begin
transpiring. As you can imagine, when this happens year after year, the large central
portion of the trunk (the heartwood) becomes wholly non-functional for the purposes of
conduction. The functioning xylem (the sapwood) is limited to the peripheral ring of the
trunk. For this reason, these trees are termed "ring-porous" (Ellmore & Ewers 1986;
Zimmerman 1983; White 1992).

At the other extreme are trees with extremely small vessels such as sugar maple (Acer
saccharum), tuliptree (Liriodendron tulipifera), and American beech (Fagus grandifolia) (see
Figure 3). The water in the vessels of these trees also freezes, but their small diameter
allows only minute bubbles to form. Most of these small bubbles are easily reabsorbed in
the spring because of the very same positive pressure that allows us to tap for maple syrup
(Zimmerman 1963). These trees can, therefore, recycle their vessels from year to year,
meaning that their functioning sapwood is large relative to their heartwood. To this recycled
sapwood, they add a fresh layer of xylem annually. Since their functioning sapwood is
spread evenly throughout the diameter of the trunk, these trees are termed "diffuseporous."
The final class of trees, which includes conifers such as white pine (Pinus strobus) and red
cedar (Juniperus virginiana), is one in which continuous vessels are wholly absent. As
depicted in Figure 3, the vessel elements of these "non-porous" trees are actually spindleshaped cells that are arranged so that their porous surfaces are aligned with each other.
These trees most likely conduct water much like diffuse-porous trees in that they use
multiple rings of xylem tissue for transportation. The main reason for this is that only tiny
embolisms can form in their vessel cells, and these can be easily reabsorbed in the spring.
Embolisms are not the only challenge facing trees. If we return to Poiseuille's law, we see
that the ring-porous trees, because of their large vessels, are much more efficient than
diffuse-porous trees. However, the increased efficiency incurs two major costs
(Zimmermann 1983). It takes greater tension to pull water up the larger vessels. This
increased tension raises the likelihood of cavitation (collapse and rupture of the vessel),
which permits embolisms to form, thereby destroying the vessel irreversibly. In addition,
the wider vessels are more susceptible to fungal infections such as the notorious Dutch elm
disease. It is thought that these are the two most salient constraints on the diameter of
vessels.
Modifying the Pipe Model
This difference in vessel conductivity between ring-porous and diffuse-porous trees means
that the model proposed by Shinozaki et al (1964a) cannot accurately portray reality for all
trees. In the wake of this realization, biologists have developed allometric models that take
these different conductivities into account (Dvorak et al 1996; Gilmore et al 1996).
Zimmermann (1978) first developed the concept of Leaf Specific Conductivity (LSC) to
quantify the relationship between the volume rate of flow through a cross-section of xylem
and the unit mass of leaves supplied by it. This differs from Shinozaki et al's model in that it
takes into account the differential conductivity of xylem rather than assuming that all xylem
conducts equally. Subsequent research has found that there is a large difference between
the LSC of diffuse-porous and ring-porous trees (Zimmermann 1983; Chapman & Gower
1991). In particular, ring-porous trees, with their large vessels, have higher LSC values,
meaning that they conduct more efficiently.
Even within individual trees, LSC differs for various parts of the tree (Zimmermann
1978).Specifically, the conductivity of the main trunk tends to stay constant while that of
the branches decreases as we follow them from their junction with the main trunk out to
their twig tips. This gradient facilitates an even distribution of water to all parts of the tree.
Some describe this phenomenon through the analogy of an open basin in which all the twigs
of a tree are dipped (Tyree & Ewers 1991; Yang & Tyree 1992). Branch junctions also tend
to have a very low conductivity. These "junction constrictions" are thought to be protective
measures to prevent excessive water loss from broken limbs (Zimmermann 1978).

The LSC correlation was modified further when it was found that not all sapwood is
conductive, especially in ring-porous trees (White 1992; Chapman & Gower 1991). White
discovered that the best correlation was to be found between the leaves and what he called
"current sapwood area" (CSA), which amounts to the growth rings of the latest year and a
half.

Putting Leonardo to the Test

These burgeoning discoveries about the hydraulic architecture
of trees have led to the current study, which examines
Leonardo's area-preserving hypothesis while considering
differences in conductivity within individual trees and among
various species of trees. Since all of the evidence cited so far
indicates that the hydraulics of a tree impacts greatly on its
architecture, it seems reasonable to say that Leonardo's areapreserving rule should also be contingent upon hydraulics. In
particular, I expect to observe two pattern;one within
individual trees and the other among species with differing
vessel structures.
Within a given tree, I anticipate a slight downward taper in
total area as I move from the main trunk to the twig tips. This
trend is predicted by the original Pipe Model (Shinozaki et al
1964a). The vessels leading to branches that have been
pruned or broken become heartwood prematurely, thus adding
to the girth below a node without actually connecting to any
leaves (see Figure 4). This means that the girth below every
node should be slightly greater than the girth above the node, even in an area-preserving
situation (Nygren et al 1993). This tapering effect could be compounded by the vessel
constrictions mentioned earlier, for in order to mitigate the risk of cavitation in distal
branches, each individual vessel tapers (Zimmermann & Potter 1982). I expect that this
taper will manifest itself in the gross structure of the tree as well.
When I compare among various species, I expect to see differences that correspond to
differences in vessel structure. Namely, I anticipate that the twigs of trees with thick piths
like black walnut (Juglans nigra) and white pine (Pinus strobus) will overshoot the areapreserving rule because the pith comprises a major proportion of the cross-sectional area.
The conductive xylem on the other hand comprises a small fraction of this cross-section. By
extension, I expect trees with little or no pith to fall closer to the area-preserving rule. As I
move to larger branches, I expect the split to be based predominantly on vessel size.
Diffuse-porous trees--sugar maple, American beech, and tuliptree--and non-porous trees-white pine and eastern red cedar (Juniperus virginiana)--should preserve area across
branchings because much of their wood is conductive sapwood. Conversely, I expect that
ring-porous trees--namely oaks, American elm, and black walnut;will preserve
circumference rather than area, since their conductive xylem is limited to the outer rings.
According to this scheme, the only trees that ought to follow Leonardo's rule for all branch
sizes are those that are both diffuse-porous and have no pith. A species with a large pith
should grow twigs that appear above the curve and should grow large branches that settle
into either the area-preserving pattern or the circumference-preserving pattern, depending

upon whether it is diffuse-porous or ring-porous, respectively. The rest of this paper is full
of surprises!

Where the Trees Grow

The breathtaking woods surrounding Princeton University in Princeton, New Jersey, served
as my laboratory throughout this study. Many measurements were taken from trees in the
woods of the Institute for Advanced Study and the Stony Ford Center for Ecological Studies.
The more elusive American elm was observed along the walkways of the university itself,
under the gaze of curious onlookers. Throughout the project, I discovered my primate roots
as I climbed many a tree.

Gathering Evidence
Fancy Equipment
For the most part, I gathered data using a simple tape measure and a caliper. I measured
trees as high as I could reach them, whether by stretching or climbing. I found that recently
fallen trees were a blessing in disguise, for I could take a myriad of measurements from the
base of the trunk to the very tips of twigs in no time at all. But waiting for trees to fall is
somewhat tedious. Fortunately, my labors were eventually eased when I was bestowed with
a hand-crafted, home-made, top-of-the-line pentaprism caliper. With this optical caliper, I
could measure trees quite accurately from a distance, without waiting for them to fall and
without regressing to apish antics.)
My Subjects
I observed ten species of trees for this study. Since I wanted to compare species with
different vessel structures, I chose several trees from each class (i.e. ring-porous, diffuseporous, and non-porous). I also chose two species that have thick piths so that I could
compare them to those that lack such a pith. Table 1 summarizes this information.

Common
name

Scientific name

Vessel
structure
Ring-porous

Presence of
pith

White Oak

Quercus alba

None

Red Oak

Quercus rubra

Ring-porous

None

Black Oak

Quercus velutina

Ring-porous

None

American Elm Ulmus americana

Ring-porous

None

Black Walnut

Juglans nigra

Diffuseporous

Present

Sugar Maple

Acer saccharum

Diffuseporous

None

American

Fagus grandifolia

Diffuse-

None

Beech

porous

Tuliptree

Liriodendron
tulipifera

Diffuseporous

None

White Pine

Pinus strobus

Non-porous

Present

Red Cedar

Juniperus
virginiana

Non-porous

None

Table 1. A summary of the ten species that are included in this study along with there vessel structures and an indication
of the presence or absence of a pith.

Measurements
I limited my measurements to even bifurcations where it was clear that one branch had split
into two. It was important to avoid taking measurements at junctions where an old branch
had broken off or had been pruned away (refer back to Figure 4 for explanation). It was
also important to take my measurements at a reasonable distance from the junction itself
because there was generally a considerable swelling above and below each junction. I found
it safe to take these measurements at a distance from the node where the swelling had
tapered down but before the swelling for the next junction began.
At each branch junction, I took three measurements: the diameter of the stem below the
node and the diameter of each of the ramifications of that stem (see Figure 5). Using these
data, I was able to analyze the branching structure according to the scheme laid out in the
following section.
Application of Trigonometry
A sensitive test was developed to analyze tree branches to determine whether or not they
follow Leonardo's area-preserving rule (Horn, in press 1998). This test uses trigonometry to
describe the relationship between the radii above and below the node. I label the radius of
the stem below the node c, and the two radii of stems above the node, a and b (see Figure
5).
The area-preserving rule asserts that the total area above the node equals the total area
below the node. In other words, (pa2+(pb2=(pc2, which is equivalent toa2+b2=c2.
The circumference-preserving rule, on the other hand, asserts that the total
circumference below the node equals the total circumference above the node. In other
words, 2(pa+2(pb=2(pc, which is equivalent toa+b=c. One final scenario is one in which
both a and b are the same as c. That is, a=b=c (I call it the doubling rule). While such a
branching pattern is unrealistic for large branches, it may be possible for small ones that
have relatively large piths.

If we now conceptualize the

radiia, b, and c as the sides
of a triangle (see Figure 6),
we find that the areapreserving rule recalls the
Pythagorean Theorem with
legs a and b and
hypotenusec. The
circumference-preserving rule
describes a distorted triangle
where the angle between
sides aand b is 180o (i.e. a
straight line). Finally, the
doubling rule produces an
equilateral triangle, where all
three sides are equal. To
determine which rule a
branch follows, we can look at
the cosine of the angle
between sides a and b(i.e.
Cos of angle C). This angle
can be described by the
following equation:

Cos (C)=(a2+b2-c2)/2ab
According to the various rules, these are the values of Cos (C):
Area-preserving rule Cos (C) = Cos (90o) = 0
Circumference-preserving rule Cos (C) = Cos (180o) = -1
Doubling rule Cos (C) = Cos (60o) = 0.5
By graphing Cos (C) against the radius or diameter of the stem below the node, I will be
able to determine which rule various sized branches tend to follow.

My Findings

All ten species that I observed followed a nearly identical pattern for branching allometry
throughout the tree. There was a tendency for twigs to follow the doubling rule, where Cos
(C) = 0.5. This trend was not limited to the pithy white pine and black walnut, as I had
predicted. Rather, it describes the trees with the thinnest twigs, American beech (see Figure
7), as well as the thickest, black walnut (see Figure 8).

Figure 7. Even though American beech has no pith, it demonstrates the same pattern as black walnut, where the twigs
tend towards the doubling rule (Cos of angle C = 0.5), while thicker branches tend towards the area-preserving rule.

Figure 8. I expected black walnut to show a unique pattern because of its thick pith. While its twigs do tend towards the
doubling rule (Cos of angle C = 0.5), black walnut is not unique in this respect (Horn, personal communication, also in
press, 1998).

For branches greater than 5 cm in diameter, the scatter decreases and a constant trend
emerges. For white oak, this trend is precisely area-preserving (see Figure 9). For all other
species, the trend is slightly above the area-preserving line, implying that cross-sectional
area is actually increasing gradually across every branching from the trunk to the twig tips
(see Appendix A for graphs of all ten species and Appendix B for a complete listing of my
data).

Figure 9. White oak shows the strictest adherence to the area-preserving rule of all the species studied (Horn, personal
communication, also in press, 1998).

Putting it All Together

The Pipe Model Falls Short
A purely hydraulic explanation of tree allometry is not sufficient to explain the trends
observed in this study. If, as I had hypothesized, vessel architecture played a significant
role in dictating allometry, the outcome would have looked very different. In particular, the
trees would have segregated into two groups based on different vessel structures and
arrangements. Data from diffuse-porous trees would have fallen along the area-preserving
line, while ring-porous trees would have dropped below this line toward the circumferencepreserving line. In addition, the twigs of trees with thick piths would have overshot the
area-preserving line and reached toward the line of the doubling rule.
If vessel architecture had played a larger role, I would also have found widespread
scattering of the data within each species' data set. I expected this scatter because the data
were collected from conspecifics in wide-ranging habitats. As I asserted earlier, tree growth
is highly dependent on the environmental conditions. For example, a tree that grows in the
midst of a dense forest would tend to grow in height faster than in width because it must
quickly reach the light in the canopy if it is to survive. By extension, an open-grown tree will
have the luxury of growing in more even proportions, since it is not constrained by light.
There was no evidence of such a difference in my observations.
I expected an even stronger confounding effect to stem from the different abilities of these
trees to cool themselves when subjected to direct sunlight. Trees with large vessels are
severely limited in their cooling capabilities because rapid transpiration would likely cause
very high negative pressures in the xylem, thereby increasing the risk of cavitation
(Zimmermann 1983). If this is true, than oaks and elms grown in full sun might be less
healthy than their forest-grown brethren. The lack of any such variation nudges me to look
elsewhere for explanations.
Interpretations

I shall begin by looking at the trends that do exist to see what they tell us about my trees
and trees in general. According to my findings, we can roughly approximate the aboveground biomass of a tree by using very simple measurements and by assuming that the
tree space is a cylinder (as illustrated in Figure 2). This approximation will work especially
well for white oaks, which appear to preserve area ideally.

Figure 10. This conical model is similar to the cylindrical model in Figure 2. However, this model may describe tree form
more accurately or at least give us an upper estimate of the volume. Note that the extreme taper of the cone is an
exaggeration.

For all the other species I examined, the cylindrical model will be an underestimate of the
actual volume, since their total cross-sectional area actually increases from the base of the
trunk to the twig tips rather than remaining constant. For these trees, the shape of the treespace can be better described by a cone (I shall call it a pseudo-cone) whose point is buried
under the ground and whose mouth flares upward toward the sky (see Figure 10) (Horn, in
press 1998).
We can attempt to calculate the volume of this pseudo-cone using the insights gained from
this study. The geometry is a little more complicated than it was for the simple cylinder, so I
will relegate the details of that calculation to an appendix (see Appendix C). This conical
model is most certainly an improved estimation of the volume of a tree, but it also has its
limitations. One of these limitations lies in the fact that most trees have a relatively lengthy
trunk that does not branch until it reaches the crown. The failure of the trunk to branch
along much of its length means that its cross-sectional area is not increasing constantly as
the conical model predicts. On the contrary, the trunk tapers from its base to the first
branching (McMahon 1975). For this reason, the conical model overestimates the volume of
the trunk itself, and by extension, overestimates the volume of the entire tree.
While both the cylindrical model and the conical model of tree form are estimates for the
volume of trees, they each serve a valuable purpose. The cylindrical model provides us with
a lower bound for our estimation of the volume. Using this value, we can approximate the
lower limit of carbon storage for a stand of woods very easily. The conical model, on the
other hand provides an upper limit for the same measurement. Even if these calculations
seem more complicated than the simple cylinder, it is still a tremendous improvement over
the alternative, which involves murdering the tree to weigh its dismembered parts. I
anticipate that further study will enable us to extract the precise shape of this cone so that
we can narrow the margin between the upper and lower bounds. The end result will be
more accurate estimations of the volume of trees than those calculated by Nygren et al
(1993).

In addition to the steady expansion in cross-sectional area that the lower parts of each tree
exhibited, the twig tips consistently expanded at an extraordinary rate, many times
approaching the line for the doubling rule. The fact that all species showed an identical
trend in the expansion of total cross-sectional area at the twig tips indicates that the pith
has no significant impact on allometry, contrary to my hypothesis. On the other hand, this
expansion in twigs does corroborate White's conjecture (1993) that it may be necessary to
have a minimum quantity of wood for a twig to be viable. This is plausible if we assume that
the leaves' demand for water is proportional to their area. The leaves supported by a
particular first-year twig are supplied by only a single growth ring. After this first year,
however, the previous year's rings can be recruited to conduct as welleven in ring-porous
species (White 1993; Ellmore & Ewers 1986). Therefore, after the first year, it suffices to
produce slightly less sapwood, while the first ring must be relatively large in order to sustain
the foliage.
A New Direction
The most peculiar result of this study is the finding that there is no detectable difference
among the allometries of trees with disparate vessel architecture. The lack of corroborating
evidence goads me to look to other possible explanations for the pervasive patterns I found.
Such an explanation may lie in a structural model of tree design (McMahon 1975; McMahon
& Kronauer 1975). McMahon proposes a model of a tree in which the main consideration is
the structural support of the individual branches and of the tree as a whole. This model
shows that trees seem to increase their girth in response to mechanical stresses caused by
bending under their own weight. In other words, increased width is a protection from
buckling. But it is not clear from their model whether this necessarily implies that area
should be preserved throughout the tree. Further inquiry into McMahon's "axe-head" model
is needed.

A Summary
From the results of this study, it appears that the dimensions of the ten species I observed
nearly obey Leonardo's rule and preserve area across every branching. The mystery that
remains is the explanation for this phenomenon. The vascular explanations that have been
presented are not adequate to explain my findings, so we must look to more universal
ecological pressures to which the trees could be responding. A structural approach looks
promising, however, the mechanics of wood are not yet understood clearly enough to
realize this goal.

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Appendices
APPENDIX A
Graphs of Ten Species of Trees Plotting Cos (C) Against the Diameter Below the Node (cm)
The following 9 figures show the results of my study for the ten trees I examined. The
graphs are arranged roughly according to the size of their vessel elements (largest to
smallest). Each graph plots the Cos of angle C (this index is described in the "Application of
Trigonometry" subsection) against the diameter of the branch below each node. All ten
species show a similar trend where thin twigs follow the doubling rule while larger branches

level off just above the line where Cos (C)=0.

APPENDIX B
Measured diameters (cm) and calculated Cos (C)
Appendix B-1
Black and Red Oak (Quercus velutina and Q. rubra)Sheet1!R1C1:R61C5
Species
Dc (cm)
Da (cm)
Db (cm)
Cos C
BO

27.22

14.48

21.33

-0.12

BO

13.69

11.59

9.71

0.18

BO

11.01

9.26

6.27

0.03

BO

9.55

5.51

8.50

0.12

BO

6.91

5.54

5.35

0.20

BO

6.65

5.00

5.22

0.15

RO

59.30

41.80

41.40

-0.02

RO

54.60

40.50

37.80

0.03

RO

49.30

32.50

40.20

0.09

RO

47.70

34.50

31.30

-0.05

RO

38.80

30.50

17.70

-0.24

RO

36.10

31.40

21.70

0.11

RO

33.70

29.30

23.20

0.19

RO

33.00

25.10

20.10

-0.05

RO

31.80

25.00

23.60

0.14

RO

26.70

21.30

12.50

-0.19

RO

25.10

18.80

13.40

-0.19

RO

16.20

13.00

10.80

0.08

RO

14.30

11.80

8.90

0.07

RO

12.22

6.05

10.95

0.05

RO

10.89

8.91

7.07

0.09

RO

10.80

6.90

8.90

0.08

RO

8.66

5.22

7.70

0.14

RO

7.70

6.11

5.67

0.15

RO

7.60

6.00

5.70

0.16

RO

7.60

5.10

6.50

0.16

RO

7.45

4.55

6.02

0.03

RO

7.32

4.30

5.86

-0.02

RO

7.32

4.87

6.53

0.20

RO

7.30

4.30

5.60

-0.07

RO

7.13

4.52

5.98

0.10

RO

6.90

4.60

5.90

0.15

RO

6.53

5.06

5.22

0.19

RO

6.50

4.90

5.40

0.21

RO

6.40

4.60

4.80

0.07

RO

6.30

4.74

4.77

0.12

RO

5.80

4.60

4.10

0.11

RO

5.79

4.58

4.17

0.13

RO

5.19

3.57

4.20

0.11

RO

4.65

3.92

3.18

0.16

RO

4.50

2.90

3.80

0.12

RO

4.04

3.53

2.90

0.22

RO

4.00

2.90

3.30

0.17

RO

3.28

2.32

2.67

0.14

RO

3.15

2.26

2.51

0.13

RO

3.10

2.10

2.50

0.10

RO

3.02

2.32

2.39

0.18

RO

2.90

2.10

2.40

0.17

RO

2.40

1.60

1.90

0.07

RO

2.30

1.85

1.65

0.14

RO

2.20

1.91

1.88

0.33

RO

2.00

1.60

1.50

0.17

RO

1.40

1.10

1.00

0.11

RO

1.30

1.00

1.00

0.16

RO

1.20

1.00

0.95

0.24

RO

0.90

0.70

0.70

0.17

RO

0.90

0.65

0.60

-0.04

RO

0.70

0.50

0.40

-0.20

RO

0.55

0.40

0.50

0.27

RO

0.48

0.30

0.34

-0.12

Appendix B-2
White Oak (Quercus alba)
(Source: Horn, personal communication, also in press, 1998)Sheet2!R1C1:R23C5
Species
Dc (cm)
Da (cm)
Db (cm)
Cos C
WO

0.30

0.20

0.25

0.13

WO

0.55

0.25

0.40

-0.40

WO

0.75

0.65

0.40

0.04

WO

0.75

0.50

0.40

-0.38

WO

1.10

0.80

0.70

-0.07

WO

1.40

0.80

1.00

-0.20

WO

2.00

1.40

1.40

-0.02

WO

2.80

2.00

1.80

-0.08

WO

4.90

3.80

3.20

0.03

WO

5.10

4.00

2.60

-0.16

WO

11.40

6.30

9.00

-0.08

WO

13.60

11.10

7.90

0.00

WO

14.90

10.20

11.10

0.02

WO

15.10

10.60

10.80

0.00

WO

17.00

8.50

15.20

0.06

WO

17.70

14.60

9.70

-0.02

WO

23.00

17.50

14.40

-0.03

WO

26.30

20.00

18.70

0.08

WO

30.40

17.10

26.30

0.07

WO

40.80

25.80

29.20

-0.10

WO

46.00

33.30

32.70

0.03

WO

53.50

48.50

0.10

27.70
Appendix B-3

Black Walnut (Juglans nigra)

(Source: Horn, personal communication, also in press, 1998)Sheet3!R1C1:R33C5
Species
Dc (cm)
Da (cm)
Db (cm)
Cos C
BW

0.40

0.30

0.40

0.38

BW

0.60

0.50

0.50

0.28

BW

0.60

0.55

0.45

0.29

BW

0.80

0.60

0.60

0.11

BW

0.90

0.80

0.80

0.37

BW

1.10

0.80

0.80

0.05

BW

1.20

1.00

1.00

0.28

BW

1.25

1.80

1.35

0.72

BW

1.25

0.80

1.10

0.16

BW

1.30

1.10

1.00

0.24

BW

1.30

0.90

0.90

-0.04

BW

1.90

1.50

1.50

0.20

BW

2.20

1.80

1.90

0.29

BW

2.20

1.80

1.50

0.12

BW

2.20

1.50

1.70

0.06

BW

2.50

1.80

1.35

-0.24

BW

2.90

1.90

2.10

-0.05

BW

3.00

2.10

2.20

0.03

BW

3.00

2.20

2.20

0.07

BW

3.20

2.30

2.50

0.11

BW

3.20

2.30

2.60

0.15

BW

4.00

2.90

3.20

0.14

BW

4.50

3.00

2.90

-0.16

BW

6.00

4.30

4.00

-0.04

BW

7.60

6.40

5.70

0.22

BW

9.90

9.20

6.20

0.22

BW

10.20

9.50

5.40

0.15

BW

21.00

19.10

10.80

0.10

BW

24.80

15.30

21.30

0.11

BW

29.30

23.90

17.20

0.01

BW

34.40

23.60

28.30

0.13

BW

35.30

27.10

20.10

-0.10

Appendix B-4
Sugar Maple (Acer saccharum)
(Source: Horn, personal communication, also in press, 1998).Sheet4!C1:C5
Species
Dc (cm)
Da (cm)
Db (cm)
Cos C
SM

0.23

0.20

0.12

0.03

SM

0.27

0.20

0.20

0.09

SM

0.28

0.20

0.20

0.02

SM

0.30

0.20

0.20

-0.13

SM

0.35

0.30

0.30

0.32

SM

0.35

0.23

0.24

-0.11

SM

0.40

0.30

0.30

0.11

SM

0.40

0.35

0.25

0.14

SM

0.40

0.33

0.30

0.20

SM

0.53

0.40

0.40

0.12

SM

0.55

0.50

0.45

0.33

SM

0.73

0.60

0.60

0.26

SM

0.80

0.55

0.60

0.03

SM

1.10

0.85

0.80

0.11

SM

1.15

0.90

0.80

0.09

SM

1.30

1.00

1.15

0.28

SM

1.40

1.25

0.85

0.15

SM

1.50

1.30

1.00

0.17

SM

1.60

1.40

1.00

0.14

SM

1.80

1.50

1.30

0.18

SM

2.50

1.90

1.80

0.09

SM

2.50

1.60

2.10

0.11

SM

2.50

1.90

2.00

0.18

SM

2.60

2.00

2.00

0.16

SM

3.30

2.60

2.40

0.13

SM

3.50

3.30

2.20

0.24

SM

3.60

3.10

2.10

0.08

SM

3.90

2.60

3.40

0.18

SM

4.00

2.90

3.60

0.26

SM

4.00

3.10

3.00

0.14

SM

4.30

3.20

3.20

0.10

SM

4.50

4.00

2.90

0.18

SM

4.70

3.90

2.80

0.04

SM

4.90

4.50

3.30

0.24

SM

5.10

4.10

3.50

0.11

SM

5.70

4.90

3.40

0.09

SM

5.70

5.40

4.00

0.29

SM

6.10

5.50

3.90

0.19

SM

7.00

4.90

6.00

0.19

SM

7.30

5.70

5.40

0.14

SM

7.80

6.40

4.90

0.07

SM

8.30

6.00

6.00

0.04

SM

11.30

9.20

7.00

0.05

SM

25.10

18.50

18.50

0.08

SM

30.20

16.90

25.10

0.00

Appendix B-5
Red Cedar (Juniperus virginiana)
(Source: Horn, personal communication, also in press, 1998)
Sheet5!C1:C5
Species
Dc (cm)
Da (cm)
Db (cm)
Cos C
RC

0.10

0.10

0.10

0.50

RC

0.20

0.15

0.20

0.38

RC

0.20

0.20

0.20

0.50

RC

0.23

0.20

0.18

0.27

RC

0.30

0.20

0.20

-0.13

RC

0.30

0.20

0.20

-0.13

RC

0.30

0.25

0.20

0.13

RC

0.30

0.20

0.20

-0.13

RC

0.40

0.30

0.40

0.38

RC

0.40

0.30

0.30

0.11

RC

0.45

0.30

0.30

-0.13

RC

0.50

0.40

0.40

0.22

RC

0.50

0.40

0.30

0.00

RC

0.50

0.37

0.40

0.16

RC

0.60

0.50

0.40

0.13

RC

0.60

0.50

0.50

0.28

RC

0.70

0.50

0.50

0.02

RC

0.72

0.55

0.70

0.36

RC

0.90

0.80

0.80

0.37

RC

0.90

0.60

0.60

-0.13

RC

1.00

0.80

0.70

0.12

RC

1.00

0.90

0.90

0.38

RC

1.00

0.80

0.70

0.12

RC

1.00

0.70

0.85

0.18

RC

1.10

0.90

0.70

0.07

RC

1.10

0.90

0.70

0.07

RC

1.30

1.00

1.20

0.31

RC

1.30

1.10

0.80

0.09

RC

1.40

1.10

1.00

0.11

RC

1.70

1.30

1.20

0.08

RC

1.70

1.10

1.40

0.09

RC

2.00

1.50

1.40

0.05

RC

2.70

1.80

2.30

0.15

RC

3.20

2.40

1.80

-0.14

RC

4.30

3.50

2.90

0.11

RC

4.70

3.50

3.00

-0.04

RC

5.00

3.70

3.00

-0.10

RC

6.70

3.70

5.20

-0.11

RC

8.70

5.30

7.10

0.04

RC

9.10

4.80

8.40

0.13

RC

11.80

6.20

10.20

0.03

RC

12.50

9.50

9.20

0.11

Appendix B-6
White Pine (Pinus strobus)Sheet6!R1C1:R19C5
Species
Dc (cm)
Da (cm)
Db (cm)
Cos C
WP

0.40

0.34

0.26

0.13

WP

0.50

0.50

0.40

0.40

WP

1.20

1.00

0.80

0.13

WP

1.80

1.30

1.40

0.11

WP

3.30

2.70

2.40

0.17

WP

12.16

8.88

8.24

-0.01

WP

18.34

14.14

10.98

-0.05

WP

20.66

17.38

12.74

0.08

WP

22.72

13.88

18.98

0.07

WP

26.92

20.46

20.24

0.13

WP

28.08

22.40

18.20

0.05

WP

31.80

17.60

26.22

-0.02

WP

32.54

24.42

24.82

0.13

WP

34.28

28.62

23.56

0.15

WP

39.58

29.80

30.36

0.13

WP

39.70

22.72

28.90

-0.17

WP

40.20

26.68

31.44

0.05

WP

40.40

27.62

31.80

0.08

Appendix B-7
Tuliptree (Liriodendron tulipifera)Sheet7!R1C1:R13C5
Species
Dc (cm)
Da (cm)
Db (cm)
Cos C
TU

5.44

3.94

4.42

0.16

TU

6.68

5.60

4.78

0.18

TU

8.92

7.70

7.00

0.27

TU

9.54

7.76

6.72

0.14

TU

11.20

8.02

8.84

0.12

TU

13.12

11.80

7.52

0.13

TU

18.30

12.48

15.18

0.14

TU

22.34

18.62

12.04

-0.02

TU

34.02

24.00

26.76

0.10

TU

44.70

33.00

30.00

0.00

TU

58.20

38.20

44.60

0.02

TU

60.40

40.10

53.50

0.19

Appendix B-8
American Beech (Fagus grandifolia)Sheet8!C1:C5
Species
Dc (cm)
Da (cm)
Db (cm)
Cos C
AB

1.90

1.30

1.50

0.08

AB

2.10

1.80

2.00

0.39

AB

2.40

1.80

2.30

0.33

AB

2.74

1.76

2.22

0.07

AB

3.70

2.70

3.10

0.19

AB

5.50

2.80

4.00

-0.29

AB

5.54

4.48

3.62

0.08

AB

5.64

4.88

3.46

0.12

AB

5.64

4.84

3.54

0.12

AB

6.50

3.90

5.90

0.17

AB

6.90

3.90

6.90

0.28

AB

9.08

7.70

6.18

0.16

AB

9.52

7.60

7.20

0.17

AB

10.32

6.98

7.98

0.05

AB

10.36

7.22

8.12

0.09

AB

12.10

9.40

8.66

0.10

AB

12.16

7.64

11.20

0.21

AB

12.80

7.00

12.40

0.22

AB

14.46

12.98

7.60

0.09

AB

15.60

11.46

11.72

0.09

AB

18.00

11.40

16.60

0.22

AB

18.28

14.58

12.38

0.09

AB

19.42

13.56

15.46

0.11

AB

23.90

16.10

19.40

0.10

AB

27.06

18.90

21.00

0.08

AB

30.70

20.50

22.70

-0.01

AB

32.50

27.80

21.90

0.16

AB

39.22

20.70

34.48

0.06

AB

47.52

30.26

39.40

0.09

AB

56.30

46.40

32.70

0.02

Appendix B-9
American Elm (Ulmus americana)Sheet9!R1C1:R37C5
Species
Dc (cm)
Da (cm)
Db (cm)
Cos C
AE

0.35

0.30

0.30

0.32

AE

0.45

0.40

0.35

0.29

AE

0.85

0.80

0.55

0.25

AE

1.10

0.85

0.85

0.16

AE

4.90

2.80

3.60

-0.16

AE

5.50

3.90

2.00

-0.71

AE

5.60

4.00

5.50

0.34

AE

9.10

4.90

8.80

0.22

AE

13.80

10.20

10.00

0.07

AE

14.70

10.40

8.00

-0.26

AE

16.50

8.10

15.90

0.18

AE

19.20

9.30

15.20

-0.18

AE

21.10

16.10

12.90

-0.05

AE

21.50

15.20

14.30

-0.06

AE

22.70

15.50

13.40

-0.23

AE

25.80

18.00

16.60

-0.11

AE

25.90

18.80

18.50

0.04

AE

30.00

21.10

23.00

0.08

AE

30.60

25.60

22.90

0.21

AE

31.10

25.60

19.70

0.08

AE

31.40

20.80

20.80

-0.14

AE

33.90

25.00

22.30

-0.02

AE

38.30

24.50

29.10

-0.01

AE

39.40

27.10

33.10

0.15

AE

39.90

31.20

27.70

0.09

AE

41.50

33.80

26.20

0.06

AE

41.70

31.90

29.30

0.07

AE

43.90

29.00

33.40

0.02

AE

48.90

38.10

38.20

0.18

AE

51.20

36.70

37.30

0.04

AE

51.30

33.70

35.50

-0.10

AE

54.20

36.70

44.90

0.13

AE

56.50

39.60

49.90

0.22

AE

58.40

37.30

52.40

0.19

AE

58.50

47.60

47.10

0.24

AE

59.50

46.70

43.20

0.13

APPENDIX C
Details for Calculating the Volume of the Pseudo-cone

We can find the volume of the pseudo-cone by finding the volume of the true cone that
extends into the ground (see Figure 11) and then subtracting the imaginary part of the cone
that is underground. The general formula for this difference of volumes is:
V = 1/3([ (D22)(H+h)/4 (D12)(h)/4]
= 1/3([ (D12)(A1B-1)(A2)(H+h)/4 (D12)(h)/4]
I shall break this formula down into its component parts to show its derivation (See Figure
11). I use V to mean volume of the tree; D2 is the diameter of the larger end of the cone,

hence(D22/4 is the area of the large end of the cone; D1 is the diameter of the small end
of the cone and is equivalent to DBH; H is the height of the tree; h is the height of the
imaginary cone that extends into the ground; A1 is the average ratio describing the
increase in cross-sectional area across each branching with the exception of the twigs; A2 is
the same ratio as it applies to the increase in area of twigs; and B is the branching order,
which Horn (in press 1998) describes as the number of branchings from the terminal twigs
to the trunk.

Figure 11. A depiction of the conical model of tree form with its dimensions labeled.
Finding H and D1 is trivial, so I will not describe the process here. To calculate h, we can
use similar right triangles to find that h = HD1/2(D2- D1). Since we cannot
measure D2 directly, we must express it in terms of D1. To do this we must determine
exactly how the diameter of the tree, or more precisely, how the cross-sectional area of the
tree, increases from the base to the terminal twigs. This is why I introduce the
terms A and B. We will define A1 as the average ratio of the total cross-sectional area
above every node to the total cross-sectional area below every node except at the terminal
twigs. In other words A1 = (a2+b2)/c2. A2 also equals(a2+b2)/c2, only it is specific to
the terminal twigs (to understand why I must make this distinction between twigs and
larger branches, refer back to the "Interpretations" subsection). SoA, in general, describes
the amount that the total cross-sectional area increases across each branching.
So we know that the cross-sectional area increases by a factor of A at every level of
branching. And we know that branching order (B) quantifies the exact number of branchings
for a particular species. So in general, AB gives us the ratio for the increase in crosssectional area over Bnumber of branchings. We can now focus this generalization to our
specific case.
The area of the wide end of the cone is (D22/4. Since we cannot measure D2 directly, we
must express this area in terms of D1 by using A and B. The area of the narrow end of the
pseudo-cone, (D12/4, will expand into the area of the wide end by a factor of (A1B1)(A2). The termA1B-1 takes care of the expansion from the trunk to the branchings just
before the terminal twigsa value that seems to be about constant according to my findings.
The term A2 takes care of the expansion at the twig tips, which tends to flare out more
than the rest of the tree. Taking both expansions into account, tells us that the area of the

wide end of the cone is (D22/4 = (D12(A1B-1)(A2)/4. This gives us all the information
that we need to calculate the volume of this pseudo-cone, and by implication, the upper
limit for the volume of a given tree.
I shall break this formula down into its component parts to show its derivation (See Figure
11). I use V to mean volume of the tree; D2 is the diameter of the larger end of the cone,
hence (D22/4 is the area of the large end of the cone; D1 is the diameter of the small end
of the cone and is equivalent to DBH; H is the height of the tree; h is the height of the
imaginary cone that extends into the ground; A1 is the average ratio describing the increase
in cross-sectional area across each branching with the exception of the twigs; A2 is the
same ratio as it applies to the increase in area of twigs; and B is the branching order, which
Horn (in press 1998) describes as the number of branchings from the terminal twigs to the
trunk.