Abstract

Beginning with Leonardo da Vinci's assertion that trees conserve total crosssectional 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 crosssectional 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 tyLOW-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).
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 "ringporous" (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 "diffuse-porous."
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 "nonporous" trees are actually spindle-shaped 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.
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

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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 area-preserving 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.
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. ringporous, diffuse-porous, 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

Presence of pith

White Oak

Quercus alba

Ring-porous

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

Diffuse-porous

Present

Sugar Maple

Acer saccharum

Diffuse-porous

None

American Beech

Fagus grandifolia

Diffuse-porous

None

Tuliptree

Liriodendron tulipifera

Diffuse-porous

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.
Application of Trigonometry

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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
to a2+b2=c2. Thecircumference-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.

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If we now conceptualize the radii a, b, and c as the sides of a triangle (see Figure
6), we find that the area-preserving rule recalls the Pythagorean Theorem with

legs a and b and hypotenuse c. The circumference-preserving rule describes a

distorted triangle where the angle between sides a and 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
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 areapreserving 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 circumference-preserving line. In
addition, the twigs of trees with thick piths would have overshot the areapreserving 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 above-ground 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 tree-space 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
A Summary
References
Chapman, J.W. & S.T. Gower (1991). Aboveground production and canopy
dynamics in sugar maple and red oak trees in southwestern Wisconsin. Canadian
Journal of Forest Research 21:1533-1543.
Cochard, H. & M.T. Tyree (1990). Xylem dysfunction in Quercus: vessel sizes,
tyloses, cavitation and seasonal changes in embolism. Tree Physiology 6:393407.
Dvorak, V., M. Oplustilova, & D. Janous (1996). Relation between leaf biomass
and annual ring sapwood of Norway spruce according to needle ageclass. Canadian Journal of Forest Research 26:1822-1827.
Ellmore, G.S. & F.W. Ewers (1986). Fluid flow in the outermost increment of a
ring-porous tree, Ulmus americana. American Journal of Botany 73:1771-1774.
Gilmore, D.W., R.S. Seymour, & D.A. Maguire (1996). Foliage-sapwood area
relationships for Abies balsamea in central Maine, U.S.A. Canadian Journal of
Forest Research 26:2071-2079.
Horn, H.S. (in press 1998). Twigs, trees, and the dynamics of carbon in the
landscape. In J. Brown & G. West, eds. Scaling in Biology. Santa Fe Institute and
Oxford University Press.

Kemp, M., ed. and translator, & M. Walker, translator (1989). Leonardo on
Painting. Yale University Press: New Haven, CT.
McMahon, T.A. (1975). The mechanical design of trees. Scientific
American 233:92-101.
McMahon, T.A. & R.E. Kronauer (1975). Tree structure: designing the principle of
mechanical design. Journal of Theoretical Biology 59:443-466.
Nygren, P., S. Rebottaro, & R. Chavarría (1993). Application of the pipe
model theory to non-destructive estimation of leaf biomass and leaf area of
pruned agroforestry trees. Agroforestry Systems 23:63-77.
Rennols, K. (1994). Pipe-model theory of stem-profile development. Forest
Ecology and Management 69:41-55.
Richter, J.P., ed. (1939). In The Literary Works of Leonardo da Vinci, vol. 1. Oxford
University Press: London.
Robichaud, E. & I.R. Methven (1992). The applicability of the pipe model theory
for the prediction of foliage biomass in trees from natural, untreated black spruce
stands. Canadian Journal of Forest Research 22:1118-1123.
Shinozaki, K., K. Yoda, K. Hozumi, & T. Kira (1964a). A quantitative analysis of
plant formthe pipe model theory. I. Basic analysis. Japanese Journal of
Ecology 14(3):97-105.
Shinozaki, K., K. Yoda, K. Hozumi, & T. Kira (1964b). A quantitative analysis of
plant formthe pipe model theory. II. Further evidence of the theory and its
application in forest ecology. Japanese Journal of Ecology 14(3):133-139.
Sperry, J.S., K.L. Nichols, J.E. Sullivan, & S.E. Eastlack (1994). Xylem embolism in
ring-porous, diffuse-porous, and coniferous trees or northern Utah and interior
Alaska. Ecology75(6):1736-1752.
Tyree, M.T. & J.D. Alexander (1993). Hydraulic conductivity of branch junctions in
three temperate tree species. Trees 7:156-159.
Tyree, M.T. & F.W. Ewers (1991). The hydraulic architecture of trees and other
woody plants. New Phytology 119: 345-360.
White, D.A. (1992). Relationships between foliar number and the cross-sectional
areas of sapwood and annual rings in red oak (Quercus rubra) crowns. Canadian
Journal of Forest Research 23:1245-1251.
Yang, S. & M.T. Tyree (1993). Hydraulic resistance in Acer saccharum and its
influence on leaf water potential and transpiration. Tree Physiology 12:231-242.
Zimmerman, M.H. (1963). How sap moves in trees. Scientific American (reprint)
March, 1963.

Zimmerman, M.H. (1978). Hydraulic architecture of some diffuse-porous

trees. Canadian Journal of Botany 56:2286-2295.
Zimmermann, M.H. (1983). Xylem Structure and the Ascent of Sap. SpringerVerlag: Berlin.
Zimmermann, M.H. & C.L. Brown (1974). Trees: Structure and Function. SpringerVerlag: New York.
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.

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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

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

27.70

48.50

0.10

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

Appendix B-3

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

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

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

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

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

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 crosssectional 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 D2directly, 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 defineA1 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). So A, 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 cross-sectional area over B number 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 (A1B-1)(A2).The term A1B-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
termA2 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 pseudocone, 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.

Former JYI staff members have gone on to win Rhodes, Marshall, and Fulbright
Scholarships, as well as NSF Graduate Research Fellowships and other graduate
research funding.
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