Bacterial wetwood, a water-soaked condition of wood, occurs in the trunk,
branches, and roots of many shade and ornamental trees (Table 1), but
is often not obvious in trees less than 10 years old. Wetwood is most
prevalent and causes the most damage to elms, especially older elms (including
American [Ulmus americana], Chinese [U. parvifolia], English
[U. procera], European [U. carpinifolia] Siberian [U.
pumila] slippery [U. rubra], and winged [U. alata] and
poplars).
Shade and Ornamental
Trees Grown in the Midwest Which Are Susceptible to Wetwood
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apple
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elm
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London plane
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redbud
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aspen
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dogwood
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magnolia
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Russian olive
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beech
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fir
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maple
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sour gum
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birch
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hemlock
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mountain ash
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sycamore
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boxelder
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hickory
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mulberry
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sweet gum
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butternut
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horsechestnut
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oak
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tulip tree
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cottonwood
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linden
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pine
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walnut
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crabapple
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locust, black
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poplar
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willow
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Wetwood wilt and dieback of elms are most common in the Midwest
and Great Plains. It is associated with the death of branches in
large cottonwoods and with the dieback and premature death of Lombardy
poplars, some as young as two years. Most trees die back slowly
over a period of several years or more, however, small trees may
wilt and die suddenly.
Symptoms vary with location suggesting that environment influences
disease development. Trees in the western United States show more
variability in internal disease development and much less bleeding
and symptom expression than trees in the East and Midwest.
Wetwood is normally not serious in most trees but as a chronic
disease it can contribute to a general decline in tree vigor, especially
of older trees growing under adverse conditions. Wetwood in landscape
trees is usually unimportant except for the disfiguring appearance
of light or dark streaks where liquid seeps out of cracks and wounds
and flows down the bark (Figure 1). As the bleeding occurs, the
liquid or flux flows down the trunk, wetting and soaking large areas
of bark. As it dries, a light gray to white incrustation is left
(Figure 1). If the exudation continues over a period of time, the
bark has a two-colored vertical column of brown with strips of white
at the margins. This liquid is toxic and commonly causes localized
death of the cambium at the base of a pruning cut (Figure 2) and
around trunk cracks. The liquid on the bark surface becomes contaminated
with mixed populations of many different kinds of airborne bacteria,
yeasts, and filamentous fungi that give it a slimy texture and often
a fetid odor. The gray to brown, foamlike foul liquid is called
slime flux or wetwood slime. The slime flux may prevent the healing
of wounds by retarding or preventing callus formation. Fluxing occurs
in Illinois from April to December but is most conspicuous during
the summer, ceasing during the winter.
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Figure
1. Wetwood of elm - light streak caused by liquiid seeping
out of cracks and wounds.
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Figure
2.
Fluxing sap of wetwood-affected tree - sufficiently toxic to prevent
callus formation and kill bark at base of pruning wound (IL Natural
History Survey photo).
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Slime flux is distinct from the white or "alcoholic"
flux that seeps from sites where various bacteria and other organisms
infect shallow, localized wounds in the inner bark and cambial region
made by insects, ice, wind, lightning, or lawn mowers, pruning tools,
and machinery. The frothy alcoholic flux is acidic, nearly colorless,
often gives off a pleasant fermentative odor, and only persists for
a short time in summer. Alcoholic flux is not related to wetwood. |
Figure
3.
Section of elm trunk with wetwood damage. Note dark brown area
(A) in isolated portions of annual growth rings and brown streaking
(B) in part of current-season growth ring (IL Natural History
Survey photo).
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CAUSAL ORGANISMS
In some trees, especially elms and poplars, bacteria are consistently
associated with wetwood and apparently cause it. In elms, the following
bacteria are commonly isolated from diseased wood: Enterobacter cloacae
(formerly Erwinia nimipressuralis), Enterobacter agglo-merans
(synonym Erwinia herbicola), Bacillus megaterium, Pseudomonas
fluorescens, and Klebsiella oxytoca.
Elm bark associated with bleeding wounds harbors large populations of
bacteria that may be disseminated by various means to infect other stem
or branch wounds; Enterobacter cloacae has been isolated from the native
elm bark beetle (Hylurgopinus rufipes), a vector of the Dutch elm
disease fungus. Infected pruning tools may also transmit bacteria from
wetwood-infected to healthy tissue.
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Symptoms
Within the trunk and larger limbs of elms, poplars, and other trees,
wetwood appears as a dark brown to black water-soaked area which in cross
section generally appears more or less circular. Wetwood liquid under
pressure sometimes spreads to the outer sapwood. In elms and poplars it
may cause gray-brown streaks or broken bands in one or more annual rings
(Figure 3). Most wetwood in the large branches and trunks of trees appears
confined to a discolored central zone and to the innermost sapwood free
of discoloration, but may extend almost to the cambium in wounded stems.
In young elms, from 15 to 30 years old, the central core of wetwood-affected
tissue tapers to a point just above the stem-root interface. It is also
as a dark brown discoloration in root tissues, often associated with a
wound, and extending upward toward the main stem. It does not usually
occur in grafted roots.
Bacterial wetwood in broadleaved trees is differentiated from normal
sapwood or heartwood by its visibly darkened color, a sour or rancid odor
due to an accumulation of fatty acids, a higher moisture content, increased
alkalinity (by one pH unit or more), decreased electrical resistance,
abnormally high gas pressure, a higher mineral content and specific gravity,
less oxygen and more carbon dioxide. It often contains methane gas. Liquid
from elm wetwood may have up to 11 times more calcium, magnesium, and
potassium cations than healthy sapwood.
Extreme gas pressures, up to 60 pounds per square inch (4.22 kg/cm2)
have been detected in discolored wetwood in elms, while gas pressures
up to 0.7 kg/cm2 are common. Pressures fluctuate seasonably with
the highest readings from May through September. The high gas pressures
frequently force liquid to seep from wounds and flow down the bark
(Figures 1). Gas is produced in wetwood-affected trees from the
metabolic activity of bacteria. The gas typically consists of 45
to 60 percent methane, 0 to 7 percent oxygen, 6 to 16 percent carbon
dioxide, 23 to 34 percent nitrogen, and 1 percent hydrogen. The
air in normal wood contains about 19 percent oxygen, 75 percent
nitrogen, and up to 5 percent carbon dioxide. Methane and hydrogen
are produced by certain bacteria under anaerobic conditions and
do not occur in normal wood. Enterobacter cloacae, and probably
other bacteria, ferment carbohydrates, producing carbon dioxide
which is converted to methane by further bacterial action. The nearly
anaerobic condition of wetwood prevents decay by wood-rotting fungi.
The first external sign of wetwood is usually the bubbling and
seepage ("bleeding" or fluxing) from wounded tissue in
V-shaped branch crotches, wounds made by removal of branches, injection
holes (for fungicides, insecticides, or nutrient uptake), and trunk
cracks, ribs, or beaks (Figures 1 and 3). Internal gas pressure
commonly reopens old wounds. The sour liquid is colorless to tan
as it oozes out and may contain up to 10 billion bacteria per milliliter.
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Figure
4. Foliage wilt caused by wetwood often followed by dying
bark of affected branches, especially in young elms (IL Natural
History Survey photo).
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Figure
5. Brown streaks produced by wetwood in young sapwood of elm
branches - can be confused with similar streaking caused by other
wilt diseases of elm (IL Natural History Survey photo).
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The water content of affected trees varies from near normal (in
white fir) to twice normal (in elms and cottonwoods). These differences,
together with a raised carbonate content, are sufficient to account
for the raised pH and for movement of water along an osmotic gradient
from normal wood into a wetwood zone.
Foliage in the tops of trees severely affected by bacterial wetwood
sometimes curl, scorch, droop, turn yellow, and defoliate from mid
to late summer; scattered branches die back or the entire crown
gradually declines over a period of years. Foliar wilting in elms
is generally believed to occur when quantities of wetwood liquid
are forced under pressure into the xylem vessels and thus are carried
to the crown. The leaves first curl upward along their margins,
the petioles wilt and the leaves drop off while still green or turn
a dull green-brown or bronze before they fall. Slowly wilting leaves
turn yellow or brown before dropping. In some cases, the leaves
turn yellow or a dull green-brown between the veins and along the
margins of leaves. Wilting is much more common in young trees; older
trees are most likely to develop a general decline in vigor or a
branch dieback in the upper crown (Figure 4).
In elms and poplars with wilting twigs and branches, wetwood appears
as gray-brown streaks or bands in the current-season wood and often
resembles wood stained by vascular wilt fungi such as Dutch elm
disease or Verticillium wilt. The streaks extend from the trunk
into small branches and twigs and merge to form solid brown rings
in the wood of one or more seasons (Figure 5).
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EFFECTS ON LUMBER
Figure
6. (Top) FLuxing can be stopped at pruning wounds, (center)
at cracks in branch crotches, (bottom) through cracks in bark of
trunk by boring holes into diseased wood where toxic sap and gas
have accumulated. (Center and bottom left). Hole should be slanted
upward so toxic sap will flow out through opening. Short piece of
threaded iron or plastic tubing inserted into hole far enough to
be firm., will carry toxic sap away from tree (Lenore Gray drawing).
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Awareness of bacterial wetwood is important to the lumber industry.
Abnormal color and moisture loss cause the lumber to be devalued.
The affected wood of oak and hemlock commonly cracks along or perpendicular
to the growth rings. The crushing strength and toughness of poplar
wetwood is inferior to normal wood. Such weakness is believed to
be due to enzymatic degradation by wetwood bacteria of part of the
binding substance(s) between cells. In standing trees this probably
promotes cracks caused by bending action, differential expansion
along temperature gradients, as well as the freezing and expansion
of water within the degraded part of the cell wall.
When frozen, the wet zones in wood are more rigid than normal wood.
A log with wetwood often shows no defect until the lumber sawn from
it shrinks and/or cracks during kiln drying. Even when no shrinkage
or cracks develop, wetwood dries only about one-third to one-half
as rapidly as normal wood and thus requires twice as much energy.
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