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COMPARTMENTALIZATION
OF
DECAY rORE
IN
TREES

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Forest Service Agriculture Information

U.S. Department of Agriculture Bulletin No. 405
July 1977
Watercolor illustrations by David M. Carroll, Warner, New Hampshire.

■^ U.S. GOVERNMENT PRINTING OFFICE : t977 0—231-254

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 COMPARTMENTALIZATIOIM
OF DECAY IN TREES
Alex L. Shigo
Plant Pathologist, U.S. Department of Agriculture,
Forest Service, Northeastern Forest Experiment Station,
Durham, New Hampshire
and
Harold G. Marx
Research Applications Staff Assistant, U.S. Department of Agriculture,
Forest Service, Washington, D.C.

Agriculture Information
Forest Service Bulletin No. 405
U.S. Department of Agriculture July 1977
INTRODUCTION report must be studied very care- compartmented plant. In a sense,
fully. The system is called a tree is made up of many trees;
The science of tree pathology CODIT each growth ring is a "tree."
emerged from studies on decay Compartmentalization Of Decay Each "tree" is divided into many
almost a century ago. Many of In Trees compartments. A compartment
the concepts developed then When the system is learned, it can be thought of as a "room,"
have changed little over the will act as the code for under- with side walls made up of rays
years. But, in the last few dec- standing a wide variety of and front and back walls made
ades some additional information defects on most tree species. up of cells that are the last to
on the decay process in trees The CODIT system is based on form in each growth ring. The
has been developed. This new two major points. First, a tree is top and bottom of the compart-
information has added to the a highly compartmented plant. ment is formed after wounding
basic story of decay without Second, after a tree is wounded, when the elements that transport
subtracting anything important the resulting defects are liquids plug up. The compart-
from it. The new expanded compartmentalized. ment is the least common de-
concept of decay is simply more To apply the CODIT system nominator of the tree. All the
complete. And this new, more it is necessary to understand that types of cells found in the woody
complete concept gives us a the new expanded decay concept stem of a tree will be found in
better opportunity to regulate and developed in the last few decades each compartment.
control decay. includes: 1) Successions of
When microorganisms invade
Dr. George H. Hepting made microorganisms associated with
tree stems through wounds, they
the first sound observations on discoloration and decay, and 2)
do so in successions. Bacteria,
compartmentalization of decay in Compartmentalization of dis-
nondecay fungi, and decay fungi
trees in 1935. His ideas acted colored and decayed wood
are often intimately associated
as a trigger for the work that associated with trunk wounds.
in this invasion process. When
followed. The work presented To begin with, decay of wood
microorganisms invade, they first
here is an expansion of his is a natural process caused by
surmount the chemical protec-
ideas. microorganisms, mainly fungi,
tive barriers set up by the tree
The information in this pub- that enter trees through wounds.
and then move into the tree from
lication is based on 16 years of Tree wounds are usually inflicted
compartment to compartment.
research by Dr. Shigo that in- by fire, weather, insects, birds,
volved complete dissections of small or large animals, or man The weakest walls of a compart-
approximately 10,000 trees— and his activities. These wounds ment are the tops and bottoms,
mostly deciduous hardwoods, at start the processes that can lead and the inner walls. The side
least 1,000 conifers, and 17 to decay, and decay is a major walls are fairly strong. When all
tropical species. Details of these cause of damage to trees. While these walls fall to the invading
studies have been published wood decay is most often caused microorganisms, there is another
elsewhere. by decay-causing fungi, these wall that begins to form. The
The purpose of this publication fungi are often intimately asso- wall formed by the cambium
is to show how most columns of ciated with bacteria and non- after wounding is the barrier
discolored and decayed wood decay fungi in the process. zone. This wall confines the in-
associated with trunk wounds in Decay is the breakdown or de- vasion to the wood present at the
trees are compartmentalized. A composition of dead organic time of wounding. The new
great number of confusing matter. It is also essential to "trees" or rings that continue
terms are given to a wide variety new life. to form are then protected from
of defects caused by discolora- To understand how trees react invasion unless new wounds are
tions and decays in trees. This to wounding and the associated inflicted. When new wounds are
book describes a system that defects by compartmentalizing inflicted at later times, multiple
makes it possible for forest columns of defect develop.
the defects, it is necessary to
managers to understand how reevaluate our concept of how An understanding of CODIT
most of these defects develop. a tree is constructed. A tree is will help to clarify many mis-
To understand the system, the considered here as a highly conceptions about decay.
Decay is a natural process.
The breakdown of dead organic
matter is essential for new
life.

Fungi are the major group of

microorganisms that decay wood
In many cases fungi accomplish
this only in association with
other organisms, especially
bacteria.
Wood-inhabiting microorgan-
isms enter trees through wounds.
Wounds start the processes that
can lead to decay. And decay is
a major cause of damage to
trees.

Decay
 Conifers

Sound heartwood

4 -^

Large
poorly healed
branch stubs

Central column
of discolored
heartwood

Poorly healed
wound

Wt^

High-quality Low-quality
pine developing pine developing
 ÍDeciduous hardwoods and
'tropical trees

Central column
of discolored wood

Well-Wafed
.branch stubs

Clear, sound
sapwood
No heartwood

•N
^tligh-quality * Low-quality
mapie dfevdoping developing
Some common wounding agents
are

¿^f •

'^y
//

1'

Birds
 Small animals

Insects

8
 Fire "'0^^

Large animals
 •I A tree is a
^ compartmented plant

ijrtvcÄnÄil 7-3iyi

10
 In a sense, a tree is made up
of many trees. Each growth ring
can be thought of as a separate
tree.

Diagrammatic
drawing
,.-Vtee of a tree.

(tee

"Tfe®

f >s

11
 In a diagrammatic way, here The rays define the side walls
is how a tree is constructed. of the compartments. The last
(The drawings are designed to few series of cells in the growth
give a general impression of ring define the inner walls. The
compartments in trees and they compartment has holes in the
are not intended as technical top and bottom walls because
anatomical descriptions of cell the flow of materials is main-
types and arrangements.) tained in a vertical direction.
But, one of the first events that
happens after wounding is a
plugging of this system above
and below the injury. Complete
tops and bottoms then begin to
develop on the compartments.
(The term "walls" is used here
in a very loose sense only to
give the mental impression of
"rooms" or compartments in
the tree.)

Pith

Ray cells form side walls of

compartments. 3

Last cells to form in growth ring

form inner walls of compartment.
2

Top and bottom of compartment

form when vertical elements plug
after wounding. 1

Compartment
12
In wood present at the time of ducting elements are plugged. Wall 3 is discontinuous be-
wounding, the tops and bottoms The rate and degree of pit closure cause sheets of ray cells are not
of the compartments are the and element plugging depend on continuous radially and longi-
weakest walls. These we will call many factors. Completion of tudinally throughout the tree.
Wall 1. The inner walls are the Wall 1 is the result of a dynamic
second in weakness—Wall 2. The process. The plugging will then Wall 4 is a much stronger,
side walls are fairly strong- set the limits for the vertical more localized version of Wall 2.
Wall 3. The strongest wall is the extension of each compartment. The area—longitudinal, tangential
one formed by the cambium When plugging occurs rapidly, —covered by Wall 4 will depend
after wounding—Wall 4. short compartments form, but on many factors: Wound size,
when plugging is slow, long type, position, severity, time of
Wall 1 is incomplete in living compartments form. year when wounding occurs,
sapwood because the conduct- and intrinsic genetic potential
ing elements—vessels, tracheids Wall 2 is continuous around to respond to wounds.
—conduct liquids in a vertical every growth ring, and from the
direction. But as heartwood top to bottom of the tree.
forms, or after wounds are
inflicted, pits close or the con-

Wall 2 is continuous around growth

ring and from top to bottom of the
tree.

Side Walls 3 are the strongest walls.

Present at time of wounding.

Inner Walls 2 are the next weakest

walls. —^

Wall 1 is incomplete until heartwood

forms or after wounding. It is not
planar.

Top and bottom Walls 1 are the

weakest walls.

Wall 3 is discontinuous.
Sheets of ray cells may end abruptly
both radially and longitudinally

Each growth ring is made up of

many compartments. 13
 Walls 2 are continuous around the
rings and from top to bottom.

A compartment

Wall 3

Wall 2

Wall 1 is incomplete until after

wounding.,

t
Walls 3 are discontinuous inward and
up and down.

A compartment

Vascular elements plug after

wounding and complete Wall 1.

14
 Continuous Walls 2

Each growth ring is made up of

many compartments.

A tree is a highly compartmented

plant.

Continuous Walls 2

15
 ection B
Section B

Secti

Barrier zone

10-year-old wound on maple

After wounding, the cambium

forms Wall 4. This wall is called
a barrier zone. It separates wood
formed before wounding from
wood that forms after wounding.

Color key to all drawings

RED—Host response to wounding,

Barrier zone Wall 4 formed after chemical reactions to STOP
wounding invasion.
GREEN—Invasion processes by
microorganisms GO through the
barriers set up by the tree.

BLUE—Infection of dead and dying

tissues on wound surface.

BROWN—Decay processes.

16
 Section A

- Bark

Section B Bark.

1—Top and bottom walls are weakest walls 2—Inner wall 3—Ray wal
4—Barrier zone equals strongest wall
17
 Deep
experimental
drill wound

Host
response

Maple

After wounding, the tree when the walls of the compart- that the walls are not absolute
reacts. Chemical barriers de- ments begin to fall to the force in their compartmentalizing
velop around the injured tissues. of the invaders, the tops and capacities and given enough
Some wood-inhabiting micro- bottoms (Wall 1) go first, then time, even the barrier zone
organisms surmount these the inner walls (Wall 2), and (Wall 4) will fall.
barriers and begin to interact then the side walls (Wall 3). But,
with the tree. The tree exerts a most of the time the barrier zone
living protective force to keep the (Wall 4) holds, and confines the
invaders out, and the invaders invaders to the wood present at
exert a strong force to get into the time of wounding. The
the tree through the wound. figures 1, 2, 3, and 4, In addi-
When microorganisms are able tion to naming the wall, also
to get in, they move from com- indicate the relative strength of
partment to compartment. And each wall. It must be emphasized

18
 Diameter of tree
at time of wounding

Wood formed
after wounding,

10-year-old wound

Barrier zone Wall 4-

Wall—1 Weakest
—2
—3
—4 Strongest

19
 The type, severity, and posi-
tion of the wound will affect the
extent of compartmentalization.
Here are some of the most
typical examples.

Wood altered slightly—NOT

heartwood

Slightly
altered
wood

Discoloration

Barrier zone Wall 4

Barrier zone
Wall 4

Red maple

20
Three severe basal wounds.
All inner compartment walls have
fallen and only the barrier zone
remains. This is how hollows
develop.

Barrier zone Wall 4

21
 A basal wound of moderate
severity. The entire "group of
trees" present at the time of
wounding was altered slightly,
and were confined by the
barrier zone. The side walls, or
rays, held here and the column
was wedge-shaped in cross-
section. The jagged vertical edge
was due to variation in confining
ability of compartment tops and
bottoms in different growth
rings.

Dark
discoloration

"%.
22
A slight basal wound. The
entire column of wood present
at the time of wounding was
altered very little, but it was
still confined by the barrier zone.
The inner walls and side walls
held firm here and only a
"moon ring" formed.

Very faint
discoloration

Diameter of tree
when wounded

Slight wound

23
 In general, the same types of
columns occur in trees that have
a heartwood core. Discolored
and decayed wood is compart-
mentalized in heartwood. When
a mechanical wound penetrates
the sapwood to the heartwood,
the column develops most
rapidly along the sapwood-
heartwood boundary.

Heartwood Heartwood

Sapwood Sapwood

Wound

Young red oak

24
RED—Host response to wounding
BLUE—Surface infection
GREEN—Discoloration
BROWN—Decay
YELLOW—Heartwood

Injured cambium

25
 Heartwood

Heartwood formation stalled near

wound.

Barrier zone in heartwood

26
 Discolored and decayed wood
associated with poorly healed
large branch stubs is compart-
mentalized in heartwood. Central
columns of defect in heartwood-
forming trees are often asso-
Injured cambium
ciated with poorly healed
branch stubs.

Young oak

Decay

27
 Here are the basic patterns of
compartmentalized columns of
discolored and decayed wood
associated with a single wound
and with several wounds at one
time shown on cross section
in nonheartwood-forming
trees.

-<>. y^,r„

28
Here are some basic patterns
for heartwood-forming trees.

Discolored
heartwood

Slightly
altered
heartwood

29
 Most trees receive many
wounds during their lives. Every
tree has some branches that
die. When the branches are
small and the //ounds close
rapidly, very little internal defect
njured follows. But, when large branches
cambium die and healing is slow, trouble
starts for the tree. Add to this
the injuries caused by other
types of wounds at irregular
time intervals and a pattern of
multiple columns begins to de-
velop. But again the tree com-
partmentalizes the injuries and
each column is separated
from the others.

Injured
cambium

Large
broken
branch

30
'M

E Wound

Injured cambium, new barrier zone

■ forming Injured
cambium

31
Wound Old barrier
zone Wall 4

New barrier
zone forming

Old barrier
zone

32
 30-year-olcl branch stub wounds

-year-old wound

20-yei

rier zone Wall

rier
e Wall 4

year-old wound

Barrier zone 4

imioarríK Iffi,

33
 Multiple columns are common
in trees. A minor wound may set
up a central column of discolored
Open branch stub-
wood and later large branches
die and heal slowly. The decay
associated with the branch stubs
Barrier zone Wall 4 does not penetrate the inner
column of compartmentalized
discolored wood.
Open branch stub

Barrier zone
Wall 4

Diameter
when first
wounded
New column of defect
does not move Into
old central column.

Wound

34
A central column of compart-
mentalized discolored wood may
be associated with large branch
stubs that healed slowly, but
did close before decay set in.
Stub
Additional columns of discolored
and decayed wood could develop
later, for example, from severe
mechanical wounds on the
trunk.

Wound

Barrier zone
Wall 4

35
 Here are some typical patterns
of multiple columns found in
nonheartwood-forming trees.

4 4 4 4

Oinnc«rtoU -fc-U fX^'i

36
In heartwoodforming trees the
patterns are the same, but they
are sometimes difficult to see
because they are often contained
in heartwood that is already
dark in color. Compartmentali-
zation explains how columns
of discolored and decayed wood
can be found in ring patterns
separated by sound wood.
4 4 4

37
A major obstacle in understand-
ing multiple columns of com-
partmentalized decay is the
difficulty in "seeing" them in
three dimensions.

Here are some examples:

Birch—Column of discoloration
and decay associated with
a wound developing Barrier zone
around a central column
of discoloration associated
with branch stubs.

Central column
of discolored wood

Stub

Old wound

38
Maple—Column of discoloration
and decay associated
with a wound developing
around a central column
of discoloration. The
entire column of wood
Wood altered slightly
present at the time of
wounding has been
altered slightly. When
this wood is dried, it Central column of
will be a different shade discolored wood
from the wood that
formed after wounding.

Wound

Barrier zone

39
Pine—Column of discolored and
decayed wood associated
with a wound developing
around a central column
of discolored heartwood
associated with poorly Central column of discolored
healed branch stubs. heartwood

Sapwood

Barrier zone Heartwood

Wound

I n.^.
40
Ash—Column of discolored and
decayed wood associated
with a wound developing
around a central column of
decay associated with old,
poorly healed branch stubs.
The barrier zone is the
ring shake boundary.

Barrier zone

Ring shake
along
barrier zon

Wound

41
Slightly altered columns of com-
partmentalized wood associated
with wounds in pines are some-
times difficult to see. Some
ring patterns of defects in pine
form when mechanical wounds
are inflicted on trees that already
have discolored heartwood.

Decay is most advan

heartwood-sapwood

Wound

Heartwood
formation
Heartwood
stalled near
formation stalled.
wound.

emfwmi fíí^

42
Another explanation for ring rots
in pine and related genera is
that some of the decay fungi,
such as Fomes pini, first get
established in the center of the
trunk through branch infections.
After establishment, new columns
of decay form at the boundaries
of the sapwood and heartwood.
The new columns are compart-
mentalized and appear as
rings.

! ■ Discolored
heartwood

Large poorly
healed stubs
)

]l^

■>

tD^V'/^tTt

43
 Old top

Sporophore

Jd i..;u Fomes pini

44
 Heartwood

Discolored heartwood

Sporophore of
Fomes pini

New column of defect not in

sapwood

White
pine

Spon

45
 The "holding power" of columns
of discolored wood can be shown
by wounds that penetrate the
columns. The new column that
forms remains separate from
the inner columns. When the
second wound is severe, decay
may develop in the new column.
This occurs in nonheartwood-
forming trees.
New barrier
zone Wall 4
Deep drill wound into sapwood and \
established central column of
discolored wood

Injured Individual column in each growth

cambium ring

Discolored
column

Nonheartwood-forming tree

46
 ilTl
Decay did not penetrate discolored
column.

manMiiftf f^-

47
And in heartwood-forming trees.

Deep drill wound penetrates

sapwood, heartwood, and central
column of discolored heartwood.

Injured
cambium

ffn^emji rr/x^

When new wounds penetrate old

columns of decay, the decay
process proceeds rapidly in the
new column.

48
'>'''Kurgll nji.

49
There are some patterns of column begins to develop. An
defects that on first inspection intricate pattern of multiple
do not appear to fit the concept columns is associated with
of compartmentalization. How- canker rot.
ever, they do. The most notable
are the defects associated with
canker rots. Canker rot fungi,
once established in a tree, keep
the wound open by slowly killing
some living tissues around the
wound. This process is the same
as re-wounding. And, when the
new wound is made, a new

New columns form as cambium is

killed each year.

Some fungi produce sterile fungus

material.
Diameter of trunk
when branch died

Fungus kills cambium and new

wounds form.

50
51
 Another type of defect that does
not appear to fit the concept are
ring shakes—separation of the
wood along the rings. The prob-
lem here is that the barrier
zone associated with the wounds
often forms far beyond the
visible column of defect. The
barrier zone acts as a partition
between the wood present at
the time of wounding and the
new wood formed after. When
growth stresses or other stresses
are applied, the wood sometimes
pulls apart along the barrier
zone. Most shakes are associated
with wounds, but not all wounds
result in shakes.
Ring shake zone

Slightly
altered
wood
Ring ^ ¡
shake
Slightly altered wood

- Large, old open wound

52
 And, yet another type of defect
pattern that appears not to fit
the compartmentalization concept
is the "island" pattern or streak
pattern often spoken of as
mineral streak. Wounds indeed
start these processes. A pattern
type that can be termed "palm
and fingers" develops. Near the
wound all the columns in the
individual growth ring are
coalesced to form the "palm."
As the discolored columns de-
velop vertically above and below
the wound, Walls 1, or the tops
Wound
and bottoms of the compartments
in individual growth rings, react
in different degrees to the
wound and the invading micro-
organisms. When such a "palm
and finger" column is viewed
in cross section at increasing
distances above and below the
wound, islands of defect appear.
Often, a few "fingers" will develop
far beyond all the others.

Barrier zon !
Wall 4

Wood formed
pn\ti»«il y^rifi after wounding

53
In sugar maple, for example, the
"palm and finger" pattern is
the mineral streak pattern asso-
ciated with sugar maple borer
wounds. This insect is a major
cause of mineral streak in sugar
maple.

54
55
The yellow-bellied sapsucker (see
page 7) inflicts several types of
wound patterns on many species
of trees. In one type of wound
pattern a large wound is the
result, and the "palm and
finger" defect pattern develops.
Again, the "fingers" may extend
great distances above and below
the wound. And cross sections
of trunks made through the
"fingers" will show islands of
defect, or mineral streaks.

■ : ~V:

' • ""K^nr,

56
 The "palm and finger" defect
pattern is also associated with
some other wounding patterns
made by the sapsuckers.

.^

Sapsucker
wounds

57
 Palm and finger

58
Sapsuckers will often wound one Not only will discolored streaks
area on a trunk. On paper birch of the "palm and finger" pattern
a swollen black band will form develop from such wounds, but
at these sites. the barrier zone that develops
after wounding may pull apart
from the inner rings to form a
complete ring shake which is
sometimes called cup shake or
loose heart.

vr^ ^

émciHíll 1^ imtMLmniTjz.

59
On some trees, such as eastern tkJ^...« ^,-

hemlock, the yellow-bellied

sapsucker inflicts many wounds
over large areas of the trunk.

"tuamil rUB,

^t^<i

60
The barrier zones that form after
wounding act as partitions be-
tween wood present at the time of
wounding and wood that forms
after wounding. When growth
stresses and other stresses
develop over long periods, the
wood may separate along the
barrier zones, and multiple ring
shakes form.

61
The "palm and finger" pattern
of defect and barrier zones that
form shake are commonly
associated with wounds made
by larger animals such as beavers
and porcupines (see page 9 ).
When wounds are severe, Walls
1 and 2 may fall to invading
microorganisms.

WHIfs

62
Squirrels wound young, smooth-
barked trees in several genera
(see page 8 ). The typical "palm
and finger" defect pattern re-
sults. In some species these
defects are called mineral
streaks.

' Bite wounds

>
1 •■•
1 1 —^

■
f

^^ ' '

■R^ ^ Canker
F" ^--1 ' -^W'
■f-aw^i from old
wound

63
Insects wound trees (see page
8) and small islands of defects
form. Each island of defect is
a small "palm and finger"
column. The columns usually
penetrate only to the depth of
the hole made by the insect.

■,-\\

^.

anmam

64
When many insects attack a tree
at the same time, a barrier zone
may develop around the entire
tree. All the wood present at
the time of wounding will be
slightly altered. When dried, the
wood present at the time of
wounding will be a different
shade from the wood that formed
after wounding.

65
The cambium miner is a small
larva of a type of fly that eats ^
its way down the cambial region
of some trees. The "tracks" of
the larvae are compartmentalized
as very long narrow streaks of
discolored wood within growth
rings.

M
66
On some trees the wounds are
not visible. This is common with
small basal wounds, especially
those caused by fire when the
tree was very young and now
the healed wound is hidden by
forest litter. The barrier zone still
forms around such wounds and
often results in ring shake. Also,
the inrolled callus serves as a
weak spot on the tree that may
continue to split vertically. Addi-
tional cracks may form from
the barrier zone outward into
the wood that forms after wound-
ing. When the cross section of
a trunk with such a pattern of
internal defect is viewed above
the base, a star-shaped pattern
of cracks radiating out from
the center will be seen.

67
With some basal wounds, long
internal cracks, called seams,
form at right angles to the
wounds at the points where the
callus inrolls.

68
As these cracks continue to
enlarge and get closer to the
edge of the trunk, they may
suddenly split out to the bark.
These secondary seams will be
at right angles to the main seam.
Most large seams start from old
wounds and move outward.

»rrlcân-9l^- '9?:í,

69
Seams will also form from ' -cir
mechanical wounds on trunks.
Once a seam begins to form, í^V-

it will usually continue to enlarge

throughout the life of the tree.
î\%

'^:r%

70
71
In summary, the compartmental- the powerful decay processes
ization of defects in trees is a continue to decompose the
survival system that is effective wood. The decaying wood pro-
most of the time—not ali of the vides nourishment for a wide
time! And after the tree dies, variety of organisms.

1^ ' mf ; \f

¿I; m
ami
72
The decomposed wood provides
nourishment for new trees.

73