Wood Science and Technology 34 (2000) 183±196 Ó Springer-Verlag 2000

Anatomy and lignin distribution

of reaction wood in two Magnolia species
N. Yoshizawa, A. Inami, S. Miyake, F. Ishiguri, S. Yokota

183
Summary Anatomical features of reaction wood formed in two Magnolia
species, M. obovata Thunb. and M. kobus DC. which are considered to be among
the primitive angiosperms, were observed. In addition, the distribution of
guaiacyl and syringyl units of lignins in the cell walls of normal and reaction
wood was examined using ultraviolet (UV)- and visible light (VL)-
microspectrophotometry coupled with the Wiesner and MaÈule reactions. The two
Magnolia species formed a tension-like reaction wood without possessing the
typical gelatinous layer (G-layer) on the upper side of the inclined stem or branch,
in which a radial growth promotion occurred. Compared with the normal wood,
the reaction wood had the following anatomical features: (1) the secondary walls
of ®ber tracheids lacked the S3 layer, (2) the innermost layer of ®ber-tracheid
walls showed a small micro®bril angle, a fact being similar to the orientation of
the micro®bril angle of the G-layer in tension wood, and (3) the amounts of lignin
decreased in the cell walls of ®ber tracheids, especially with great decrease in
proportion of guaiacyl units in lignins. In addition, VL-microspectrophotometry
coupled with the Wiesner and MaÈule reactions adopted in the present study
showed potential to estimate the lignin contents in the cell walls and the
proportion of guaiacyl and syringyl units in lignins.

Introduction
Woody plants develop special tissues which are called reaction wood when their
stems are bent. Reaction wood is formed to return the axis of a leaning tree to the
normal or vertical position, or to maintain the natural equilibrium position with
regard to branches (Onaka 1949; Westing 1965; Timell 1986). In angiosperms,
radial growth increases on the upper side of an inclined stem or branches, and
reaction wood referred to as tension wood generally develops on this side.
Tension wood can be characterized by the presence of gelatinous (G-) ®bers
which contain a thickened cellulosic inner layer (Onaka 1949; Dadswell and

Received: 15 July 1998

N. Yoshizawa (&), A. Inami, S. Miyake, F. Ishiguri, S. Yokota
Laboratory of Forest Products, Department of Forest Science,
Faculty of Agriculture, Utsunomlya University, Utsunomlya,
321±8505, Japan

The Authors are grateful to Dr. T. E. Timell for his valuable

suggestions and providing the literature, and to Dr. T. Fujil,
Forestry and Forest Products Research Institute, Tukuba, Japan,
for his help in measuring the UV-absorption spectra.
 Wardrop 1955; CoÃte and Day 1965; Araki et al. 1982, 1983). Okuyama et al. (1990,
1994) revealed the important role of micro®brils (G-layer) in the generation of
growth stresses in tension wood of hardwood species. On the other hand, some
species of primitive angiosperms have been reported to frequently lack G-®bers
(Onaka 1949; HoÈster and Liese 1966; Kucera and Philipson 1977a, b, 1978; Maylan
1981; Timell 1986; Yoshizawa et al. 1993a, b). Members of the genus Magnolia
which is considered to be the primitive angiosperm also seem to lack G-®bers in
the reaction wood (Onaka 1949; Okuyama et al. 1990, 1994). However, no detailed
information is available about the ultrastructure and chemical composition of
reaction wood ®bers in Magnolia species.
184 Tension wood differs from normal wood in chemical composition. Timell
(1965) pointed out that abnormally low lignin contents typify hardwood reaction
wood. This fact also holds for species which do not produce G-®bers (CoÃte and
Day 1965). In hardwood species, the secondary walls of vessels generally contain
predominantly guaiacyl lignin, whereas those of ®bers contain a high proportion
of syringyl lignin (Fergus and Goring 1970a, b; Musha and Goring 1975; Saka
et al. 1983; Fujil et al. 1987). However, the formation and distribution of struc-
tural lignin units in the cell walls of reaction wood in species which do not form
G-®bers have not yet been investigated.
In this paper, anatomical features of the reaction wood in two Magnolia spe-
cies, M. obovata Thunb. and M. kobus DC., were observed. In addition, distri-
bution of guaiacyl- and syringyl-type lignins in the cell walls of both normal and
reaction wood was studied by ultraviolet (UV)- and visible light (VL)-micro-
spectrophotometry after the Wiesner and MaÈule reactions.

Materials and methods

Straight trees, 3 m tall, of Magnolia obovata Thunb., growing in the nursery at
Utsunomiya University, Japan, were inclined at an angle of 70 degrees from the
vertical in late May 1994, and then cut down after two months from the stem
inclination. For Magnolia kobus DC., branches were collected from a 40-year-old
tree growing on the campus. Small wood specimens were obtained from xylem of
the lower (opposite wood), lateral (lateral wood: referred to as normal wood
here), and upper (reaction wood) sides of the leaning stem or branch, and then
®xed with 3% glutaraldehyde. Transverse sections, 8 lm thick, were sliced from
the specimens, and stained with safranin-light green or zinc chloride-iodine so-
lution to detect G-®bers. Some sections without staining were observed under
polarizing and ¯uorescence microscopes to ®nd the secondary wall layers of the
®ber tracheids and vessels, and the extent of ligni®cation in their secondary walls.
The cell wall structure of these elements was observed on the radial surfaces with
an SEM.
Transverse sections of 8 lm in thickness were used for measuring the VL-
absorption spectra of secondary walls in the ®bers and vessels after the Wiesner
and MaÈule color reactions. The MaÈule reaction was carried out as follows: 1) 5
minutes' immersion in 1% KMnO4 and three times washing with distilled water,
2) immersion in 3% HCl for 1 minute, followed by washing with distilled water, 3)
immersion in 29% NH4OH for 1 minute, followed by washing with distilled water.
The Wiesner reaction was performed as follows: 1) pouring a few drops of 1%
phloroglucinol ethanol solution on the section mounted on a glass-slide, 2)
adding 1 drop of 35% HC1, 3) covering the section with cover slip.
These sections were used for measuring absorption spectra from 400 to 700 nm
in 1 nm steps using a microspectrophotometer (Carl Zeiss UMSP 80: spot
diameter: 0.5 lm, band width: 5 nm). Measurements were repeated 10 times at
every step for a point measured in the secondary walls of the ®bers and vessels.
Only one section was used for each measurement of the spectra. Due to the
temporary nature of the color reactions, all measurements were performed within
10 minutes. Ten VL-absorption spectra obtained from each measurement were
averaged to give the mean spectra.
For measuring the UV-absorption spectra of the ®ber and vessel walls, trans-
verse sections, 2 lm thickness, were mounted on a quartz slide, immersed in
glycerin, then covered with a quartz cover slip. Ultraviolet absorption spectra of
these sections were measured from 260 to 300 nm in 1 nm steps according to the
procedure of Fujii et al. (1987). 185

Results and discussion

Reaction wood anatomy

Two species Magnolia tested here exhibited a radial growth promotion on the
upper side of the inclined stem or branch. The ratios of the growth increment on
the upper side/lower side were 3.8 for M. obovata and 2.3 for M. kobus, respec-
tively. Formation of G-®bers was not observed on either the upper side or lower
side of the inclined stem or branch in both species (Figs. 1 and 2). It was con-
®rmed by the histochemical observation that neither light green staining nor zinc
chloride-iodine treatment indicated the presence of a clearly de®ned cellulosic
layer (G-layer) characteristic of tension wood. Remarkable reduction in the size
and number of vessels was found in xylem formed on the upper side in both
species (Tables 1 and 2). This feature may also be common to some species which
develop particular forms of reaction wood without a distinct G-layer in their
®bers (Scur®eld and Wardrop 1962; Kucera and Philipson 1977a, b). Almost no
differences were found in the diameter and wall thickness of the ®ber tracheids
among the upper, lateral, and lower sides of the samples, whereas the cell length
slightly increased on the upper side. However, the cell wall structure of reaction
wood ®bers differed from that of the lateral and opposite woods. As shown in
Fig. 3, the secondary walls of ®ber tracheids of the lateral and opposite woods had
the usual three-layered structure (S1 + S2 + S3) in both species, whereas reaction
wood ®bers lacked the S3 layer. This fact is in agreement with the feature of
reaction wood ®bers of Buxus, considered to be one of the primitive angiosperms,
which produced compression wood on the lower side and no G-layer on either the
upper or lower side (Yoshizawa et al. 1993a, b).
In addition, it is particularly interesting to note that the innermost layer of
secondary wall layers showed a small micro®bril angle, as shown in Fig. 4. This
feature is very similar to that of the G-layer which is made of cellulose micro®brils
oriented nearly parallel to the longitudinal axis of the cell (Araki et al. 1982, 1983;
Prodhan et al. 1995a, b). This fact is in agreement with the results previously re-
ported by Okuyama et al. (1990, 1994), who showed that the secondary walls of
tension wood ®bers have small micro®bril angles (5 to 10 degrees) in M. acuminata
L. and Liriodendron tulipifera L., belonging to the same Magnoliaceae, which
produced no G-layer. However, Fig. 5 apparently demonstrated that in reaction
wood ®bers the outer part of the S2 layer has greater micro®bril angles than those of
the innermost layer. Nearly no differences in the micro®bril angle of the outer part
of the secondary wall were found between the normal and reaction wood, except for
the innermost layer. This probably indicates that the micro®bril angle of the sec-
ondary ®ber walls gradually decreases during the transition from normal wood to
186

Fig. 1. Fluoresence microphotographs of transverse sections without staining in the

normal wood (A) and reaction wood (B) formed on the lateral and upper sides, respec-
tively, of M. obovata. Formation of G-layers is not observed on the upper side of inclined
stem, and reduction in the size and number of vessels is seen. Scale bar ˆ 30 lm

Fig. 2. Fluorescence microphotographs of transverse sections without staining in the

normal wood (A) and reaction wood (B) of M. kobus. Formation of G-layers is not observed
on the upper side of branch, and reduction in the size and number of vessels is seen.
Scale bar ˆ 30 lm
Table 1. Reaction wood anatomy of M. kobus

Cell type Diameter (T) Thickness Length Distribution

(lm) (lm) (mm) (No./mm2)

Fiber tracheid RW 14.3 2.3 1.1 ±

LW 14.5 2.2 1.0 ±
OW 14.0 2.2 0.9 ±
Vessel RW 36.3 1.5 ± 46.6
LW 41.5 1.6 ± 85.6
OW 38.1 1.5 ± 82.2

RW Reaction wood, LW Lateral wood, OW Opposite wood 187

Table 2. Reaction wood anatomy of M. obovata

Cell type Diameter (T) Thickness Length Distribution

(lm) (lm) (mm) (No./mm2)

Fiber tracheid RW 15.1 2.1 1.2 ±

LW 14.9 1.9 1.0 ±
OW 14.8 2.1 0.9 ±
Vessel RW 38.8 1.4 ± 34.0
LW 45.3 1.5 ± 58.0
OW 38.3 1.5 ± 54.0

RW Reaction wood, LW Lateral wood, OW Opposite wood

reaction wood. In the species which have no G-®bers, it was revealed that (a) the
smaller the micro®bril angle, the larger the longitudinal tensile stress, and (b) the
larger the tensile stress, the larger the a-cellulose content (Okuyama et al. 1990,
1994). It is considered that the change in the micro®bril angle occurring when
normal wood is transformed into reaction wood probably corresponds to the in-
tensity of the stimulus for reaction wood formation (tensile stress): that is, the
increase in a-cellulose content suggests the increase in degree of polymerization for
cellulose, irrespective of whether a G-layer is formed or not.
Kucera, Philipson (1978) and Maylan (1981) studied reaction wood anatomy of
Pseudowintera colorata, a vessel-less dicotyledon, which exhibited a pronounced
growth promotion on the lower side of inclined branches. They also reported that
the only signi®cant difference between the anatomy of the upper and lower sides
is that the tracheids on the lower side have a larger micro®bril angle. This was
also true for Drimis wintera which developed neither compression wood nor
tension wood (Kucera and Philipson 1977a). It is very interesting to note that
there is a difference in micro®bril angle of the secondary tracheid walls between
these species which produce neither compression wood nor tension wood. The
only anatomical difference between the upper and lower sides of the inclined stem
or branch is that the side of growth promotion apparently differs among these
species. In Buxus wood a radial growth promotion occurred on the lower side of
the inclined stem, in which the secondary walls of ®ber tracheids increased the
lignin contents and the micro®bil angle (Yoshizawa et al. 1993b). These former
results and those obtained here suggest that tensile stress tends to decrease the
micro®bril angle, accompanied by the increase in a-cellulose content, in the
secondary walls of axial elements with a function of mechanical support. This is
in contrast to the fact that compressive stress increases the micro®bril angle,
188

Fig. 3. Polarizing microphotographs of transverse sections in M. obovata (A, B) and M.

Kobus (C, D). A, C ˆ normal wood, B, D ˆ reaction wood. Fiber tracheids lack S3 layer in
reaction wood of both species. Arrowheads indicate the presence of S3 layer in normal wood
®bers. Scale bar ˆ 50 lm

accompanied by the increase in lignin content. Further research is needed to

examine the changes of micro®bril angle in the secondary ®ber-tracheid wall
during the reaction wood formation of the other Magnolia species in more detail.

Staining properties with Wiesner and MaÈule reactions

As shown in Figs. 6 and 7, nearly no differences in the intensity of auto¯uores-
cence from lignins were found in the secondary walls of the ®ber tracheids and
vessels between normal and reaction wood. Transverse sections after the Wiesner
reaction demonstrated that in normal wood the vessel walls and middle lamella
are stained in deep red, whereas the ®ber walls are not deeply stained (Fig. 8). A
remarkable difference in the staining property was found in the secondary walls
of ®ber tracheids of normal and reaction wood. The secondary walls of ®ber
tracheids were less stained in reaction wood than in normal wood, whereas there
were almost no differences in the staining intensity of the secondary walls of
vessels between normal wood and reaction wood. On the other hand, the MaÈule
reaction gave almost no differences in the staining property of the secondary
walls of the ®ber tracheids and vessels between normal and reaction wood
(data not shown).
The Wiesner reagent reacts with coniferyl and sinapyl aldehyde units in lig-
nins, however, coloration may be weak or even absent in lignins containing high
amounts of syringyl units (Sarkanen and Ludwig 1971; Nakano and Meshitsuka
1992). In contrast to this fact, coloration by the Wiesner reaction becomes pro-
gressively stronger with the increase in Klason-lignin content (Sarkanen and
 189

Fig. 4. Scanning electron microphotographs of radial surfaces in the normal wood (A) and
reaction wood (B) of M. kobus. Note the lack of S3 layer in the ®ber tracheids of reaction
wood and the small micro¯bril angle of the innermost layer. Scale bar ˆ 10 lm

Hergert 1971). The results obtained here suggest that ®ber tracheids decreased
their lignin contents, particularly of guaiacyl units in lignins compared with the
syringyl units, during their changes from normal wood to reaction wood. This
fact was also con®rmed by the VL- and UV-microspectrophotometry as described
later.

VL-absorption spectra
Figures 9 and 10 show VL-absorption spectra of the ®ber-tracheid and vessel walls
after the Wiesner and MaÈule reactions in M. obovata and M. kobus. Both species
showed very similar VL-absorption spectra in normal and reaction wood. In
normal wood, an absorption maximum of around 570 nm for Wiesner reaction
and around 515 nm for MaÈule reaction was observed in the secondary walls of the
®ber tracheids and vessels, respectively.
190

Fig. 5. Scanning electron microphotographs of radial surfaces of ®ber tracheids in the

normal wood (A) and reaction wood (B) of M. kobus. In reaction wood, the outer part of the
secondary wall layer shows greater micro®bril angles than those of the inner part. Arrows
indicate the orientation of micro®brils. Scale bar ˆ 10 lm

Fig. 6. Fluorescence microphotographs of transverse sections without staining in the

normal wood (A) and reaction wood (B) of M. obovata. Strong auto¯uorescence of lignin
can be seen in the secondary walls of the vessel and middle lamella in both normal and
reaction wood. Almost no differences in the auto¯uorescence intensity of lignins between
both woods are observed in the secondary walls of the ®ber tracheids and vessels.
Scale bar ˆ 10 lm

The spectra obtained with Wiesner reaction exhibited an absorption maximum

of around 570 nm, also in reaction wood of both species. In normal wood of both
 191

Fig. 7. Fluorescence microphotographs of transverse sections without staining in the

normal wood (A) and reaction wood (B) of M. kobus. Almost no differences in the
aouto¯uorescence intensity of lignin between both woods are observed in the secondary
walls of the ®ber tracheids and vessels. Scale bar ˆ 10 lm

species, the absorbance was greater in the vessel walls than in the ®ber tracheid
walls. In reaction wood, the vessel and ®ber-tracheid walls exhibited lower ab-
sorbance than that of normal wood, considerably low for the ®ber tracheids with
absorbance ratio of about 0.6 in both species (Figs. 9 and 10). This fact is in
agreement with the results of the microscopic observation of the transverse
section after the Wiesner reaction. The greater decrease in the absorbance may
indicate that the amount of guaiacyl units in lignins decreased to a larger extent in
the secondary walls of ®ber tracheids by the formation of reaction wood, con-
sidering that the secondary walls of ®ber tracheids showed low UV-absorption
in reaction wood, as described later.
The spectra obtained with the MaÈule reaction showed appreciable absorption
around 515 nm in all tissues, indicating the presence of syringyl units (Yoshinaga
et al. 1989, 1992; Nakano and Meshitsuka 1992; Takabe et al. 1992; Yoshizawa
et al. 1993b). Although these spectra all showed a slight decrease in absorbance
around 515 nm in all tissues of reaction wood, no great decrease in the absor-
bance of ®ber-tracheid walls was observed in both species, a fact in contrast to the
results of Wiesner reaction. However, a shift of the absorption maximum position
to a shorter wavelength was observed in the ®ber walls of reaction wood (Figs. 9
and 10). In strongly MaÈule-positive tissues, the lignin is shown to be rich in
syringyl units (Srivastava 1966). Using UV-microspectrophotometry, Yoshinaga
et al. (1992) demonstrated that the position of absorption maximum shifts to a
shorter wavelength with the increase of the proportion of syringyl units in lignins
in oak wood cell walls. Fujii et al. (1987) also revealed that the UV-absorbance of
the secondary walls of ®ber becomes smaller in hardwoods which contained
smaller amounts of Klason lignin and a larger ratio of syringyl/guaiacyl (S/V)
units. What does the shift of the absorption maximum position in reaction wood
192

Fig. 8. Transverse sections after Wiesner reaction in the normal wood (A) and reaction
wood (B) of M. obovata. In normal wood, the vessel walls and middle lamella are stained
in deep red, while only slightly red the ®ber-tracheid walls. The staining intensity of ®ber-
tracheid walls is smaller in reaction wood than in normal wood. Scale bar ˆ 50 lm

found with the MaÈule reaction mean? This may be due to the change of the
chemical structure of the syringyl units in reaction wood or due to the decrease in
proportion of guaiacyl units in lignins by the formation of reaction wood. In fact,
Betula ®ber walls being rich in syringyl units showed an absorption maximum
position of around 500 nm with the MaÈule reaction, whereas Buxus ®ber tracheids
being relatively rich in guaiacyl units showed the same at around 515 nm
(Yoshizawa et al. 1993b).

UV-absorption spectra
Table 3 shows UV-absorbances in the secondary walls of the ®ber tracheids and
vessels in the normal wood and reaction wood of M. obovata and M. kobus. The
secondary walls of both elements showed an absorption maximum at 279 nm in
normal wood, whereas those of reaction wood ®bers showed it at 277 nm. There
were no remarkable differences in the UV-absorbance at 279 nm, indicating the
presence of guaiacyl units in the normal wood cell walls of both species. In
reaction wood, however, a decrease in the absorbance at 279 nm was recognized
in both ®ber tracheids and vessels, especially with greater decrease in the ®ber
tracheids. The absorbance ratio of reaction wood/normal earlywood in the ®ber-
tracheid walls was relatively small compared with that in vessels. The ratio was
0.56 for M. obovata and 0.59 for M. kobus. This fact indicates that reaction wood
largely decreased the proportion of guaiacyl units in lignins in the ®ber-tracheid
walls. These results are in agreement with those obtained by the Wiesner reaction,
suggesting that both Wiesner and MaÈule reactions have the potential to estimate
 193

Fig. 9. Visible-light absorption spectra of the ®ber-tracheid and vessel walls after the
Wiesner and MaÈule reactions in M. obovata. The spectra show absorption maximum of
around 570 nm for the Wiesner reaction, and that of around 515 nm for the MaÈule reaction
in the normal wood. The great decrease in absorbance is found in the ®ber-tracheid walls of
reaction wood in the case of Wiesner reaction. Note the shift of the absorption maximum
position to a shorter wavelength for the ¯ber-tracheid walls of reaction wood in the case of
the MaÈule reaction

lignin content in cell walls, or to re¯ect the proportion of guaiacyl units in the
lignins, as pointed out previously by Yoshizawa et al. (1993b).
Yoshizawa et al. (1993b) investigated the lignin distribution in the secondary
walls of the ®ber tracheids and vessels of Buxus microphylla which produced no
G-®bers, and revealed that both cell types increase their contents of guaiacyl
units, particularly in the outer parts of the secondary walls, during their changes
from normal wood to reaction wood. In Buxus reaction wood, the increased
proportion of guaiacyl units in the lignins obtained with the UV-microspectro-
photometry was in good agreement with the results obtained with VL-micro-
spectrophotometry coupled with the Wiesner reaction. In this paper, it is of great
interest to note that the secondary walls of ®ber tracheids have shown a similar
absorbance ratio (about 0.6) of reaction wood/normal wood with both methods
(Figs. 9 and 10, Table 3). Chemical analysis has also shown the decrease of Kla-
son-lignin content in reaction wood: decrease of 4.8% in M. obovata and of 4.3%
in M. kobus, respectively, compared with the lignin content of normal wood of
both species. This trend corresponds to the results obtained with UV- and VL-
microspectrophotometry. This means, the decrease in lignin content is consid-
ered to be mainly due to the decrease of guaiacyl units in lignins.
194

Fig. 10. Visible-light absorption spectra of the ®ber tracheid and vessel walls after the
Wiesner and MaÈule reactions in M. kobus. The great decrease in absorbance for the Wiesner
reaction can be seen in ®ber-tracheid walls of the reaction wood. The MaÈule reaction shows
the shift of the absorption maximum position to a shorter wavelength in the ®ber-tracheid
walls of reaction wood

Table 3. UV-absorbance in
the secondary walls of normal Wood type Absorbance [log(Io/I)],
and reaction wood tissues 279 nm
in M. kobus and M. obovata
Fiber tracheid Vessel

M. kobus Normal earlywood 0.37 0.42

Reaction wood 0.22 0.34
(Ratio/Earlywood) (0.59) (0.81)
M. obovata Normal earlywood 0.36 0.48
Reaction wood 0.20 0.38
(Ratio/Earlywood) (0.56) (0.79)

Absorbance was measured in the secondary walls of ®ber

tracheid and vessel at 279 nm

Conclusions
Magnolia species formed a tension-like reaction wood, though not typical
G-®bers, on the upper side of the inclined stem or branches so that the innermost
layer of ®ber-tracheid walls had a small micro®bril angle, and the amounts of
lignin in the cell walls decreased on this side, especially with great decrease of
guaiacyl units in lignins. The present study also suggested that VL-microspec-
trophotometry coupled with the Wiesner and MaÈule reactions has the potential
to estimate the lignin contents in cell walls and the proportion of guaiacyl and
syringyl units in lignins.

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