Document Text Contents
Page 2
Handbuch der Pflanzenphysiologie • Eneyelopedia of plant physiology
Gesamtdisposition:
Teil I. Allgemeine Grundlagen.
Band I. Genetische Grundlagen physio-
logischer Vorgänge. Konsti-
tution der Pflanzenzelle.
" ll. Allgemeine Physiologie der
Pflanzenzelle.
Teil ll. Stoff- und Energiewecbsel.
Band Ill. Pflanze und Wasser.
" IV. Die mineralische Ernährung der
Pflanze.
"
V. Die 002-Assimilation.
" VI. Aufbau, Speicherung, Mobili-
sierung und Umbildung der
Kohlenhydrate.
" VII. Stoffwechselphysiologie der
Fette und fettähnlicher Stoffe.
" VID. Der Stickstoff-Umsatz.
" IX. Der Stoffwechsel der schwefel-
und phosphorhaltigen Verbin-
dungen.
" X. Der Stoffwechsel sekundärer
Pflanzenstoffe.
" XI. Die Heterotrophie.
" Xll. DiePflanzenatmung, einschließ-
lich Gärungen und Säurestoff-
wechsel.
" Xlli. Der Stofftransport in der
Pflanze.
Teil ID. W acbstum, Entwicklung,
Bewegungen.
Band XIV. Wachstum und Wuchsstoffe.
"
"
XV. Differenzierung und Entwick-
lung.
XVI. Außenfaktoren in Wachstum
und Entwicklung.
" XVII. Physiologie der Bewegungen.
1. Bewegungen durch EinflüBBe
mechanischer und elektri-
scher Natur sowie durch
Strahlungen.
2. Bewegungen durch EinflÜBBe
der Temperatur, Schwer-
kraft, chemischer Faktoren
und aus inneren Ursachen.
" XVID. Sexualität, Fortpflanzung,
Generationswechsel.
Synopsis of the eontents:
Part I. General loundations.
Vol. I. Genetic control of physiological
processes. The constitution of
the plant cell.
" ll. General physiology of the plant
cell.
Part ll. Metabolism.
Vol. Ill. Water relations of plants.
" IV. Mineralnutrition of plants.
" V. The assimilation of carbon
dioxide.
" VI. Formation, storage, mobiliza-
tion, and transformation of
carbohydrates.
" Vll. The metaboliBm of fats and
related compounds.
" VIII. Nitrogen metabolism.
" IX. The metabolism of sulfur- and
phosphorus-containing com-
pounds.
" X. The metabolism of secondary
plant products.
" XI. Heterotrophy and heterotrophic
plants.
" Xll. Plant respiration, incl. fermen-
tations and acid metabolism in
plants.
" xm. Translocation in plants.
Part m. Growtb, development,
movements.
Vol. XIV. Growth and growth substances.
" XV. Differentiation and develop-
ment.
" XVI. External factors affecting
growth and development.
" XVII. Physiology of movements.
1. Movements due to mechani-
cal and eleotrical stimuli
and to radiations.
2. Movements due to the ef-
feots of temperature, gra-
vity, ohemical faotors and
internal faotors.
"XVID. Sexuality, reproduotion, alter-
nation of generations.
Page 434
418 F. F. NoRD and G. DE STEVENS: Ligninsand lignification.
C. Infrared absorption spectra.
The infrared spectrum of spruce nativeligninwas first determined by JoNES
(1949 [2]). He found it tobe generally similar to lignins isolated by strong chemi-
cal reagents. Since then the infrared spectra of the native and enzymically
liberated lignins from white Scots pine, oak, birch, maple, bagasse and "kiri"
wood have been determined by NoRD and coworke-rs. In all cases, they were
A
B
c
D
6'?5
Nolive birch Lignin
#olive ookfignin
No live mop/e Lignin
O.~~~~~~~~~LUlU~~UL~LULU~~~~LuLU~uJ
Z.O 5.0 6'.0 7.0 8.0 !l.O 10.0 11.0 1Z.O 1J.O 1'1.0 75.0 16'.0
Wove Lengtll [ Microns]
Fig. 5. Infrared absorption spectra. A White scots pine native Iignin; B birch native Iignin; C oak native Iignin;
D maple native Iignin.
found to resemble closely the spectrum of spruce native lignin. A few of these are
presented in Figure 5.
The band at 3400 cm.-1 is due to the presence of bonded hydroxyl groups in
the lignin complex. However, the intensity of the absorptionband in this region
is less pronounced in the case of bagasse native lignin than in the other lignin pre-
parations. This supports the chemical data in that bagasse native lignin contains
fewer hydroxyl groups than the other lignins. Absorption from 1700 to 1660 cm. -1
is characteristic for the presence of an aldehyde or ketone carbonyl group. This
Page 435
Electrophoresis. 419
band is very weak in the case of maple lignin. Phenyl ring skeletal vibrations with
possible para substitution are evidenced at 1600 and 1510 cm.-1• The 1430 cm.-1
band establishes the presence of aliphatic groupings in the lignin complex. A
difference in relative intensity at 1325 cm.-1 should also be pointed out. Such
variations aremoreevident at 1265, 1255 and 1220 cm.-1, which are in the spectral
region of aromatic or aliphatic unsaturated C-0 groups, and also at 1140, 1163,
1155 and 1125 cm.-1 • These differences possibly are due to different atomic
environments in the lignin complex, and since bands in this region arise from
vibrations of the molecule as a whole, interpretation cannot be specific. The
bands at 2945, 1460, 1380 and 725 cm. -1 are due to Nujol absorption, and thus are
of no interpretative value. Bagasse native lignin and kiri native lignin have sharp
bands at 835 cm. -I, whereas pronounced bands at 890 and 870 cm. -1 are found
in the maple lignin curve. JONES (1949 [3]), using a film technique (evaporation
of a dioxane solution of spruce native lignin) reported bands in the same region
even after prolonged drying. Dioxane is known to absorb in this part of the spec-
trum, but since mineral oil mulls were used with the other lignins, it can safely
be stated that the bands are characteristic of the lignin studied. These bands
are suggestive of strong absorption by trisubstituted phenyl groups. Therefore,
a comparative evaluation of the absorption spectra of these lignins reveals that
they are generally similar, but noticeable differences are observed in the relative
intensities of the bands, suggesting some deviations in the arrangement of groups
present therein. Moreover, it was found that the spectra of the enzymically
liberated lignins from each species of wood were identical with the corresponding
native lignin fractions.
D. Electrophoresis.
The literature is devoid of any studies on the electrophoretic behavior of
lignins. However, due to the presence of at least one phenolic hydroxyl group per
lignin building unit, it appeared that such an investigation would be feasible.
Moreover due to the success in applying electrophoretic methods to the characteri-
zation and fractionation of proteins, a study of the possible applicability of the
method to lignins was undertaken at Fordham.
Preliminary measurements had indicated that the native lignins from white
Scots pine and oak woods are electrophoretically inhomogeneous. Therefore,
the effect of repeated precipitation of lignin on the electrophoretic patterns
was first studied. In Figure 6 are presented the patterns obtained with the oak
lignin after four (1) and eight (2) precipitations. It can easily be recognized
that the oak lignin appears electrophoretically homogeneous, whereas the patterns
obtained with the white Scots pine lignin are characteristic of a multicomponent
system. Whereas occasionally two well defined peaks appeared, the inhomogeneity
revealed itself principally in the existence of trailing boundaries. The inhomo-
geneaus white Scots pine lignin contains components of both higher and lower
mobility than the main components.
The oak lignin was electrophoretically analyzed after every precipitation.
Under identical conditions (PR 10, KCI = 0.1 M, sodium glycinate-HCl = 0.05 M),
the electrophoretic mobility was not affected by the purification to any significant
degree. The mobility of the ascending boundary was found to be always slightly
higher than that of the descending boundary, the respective values being
-10.7 ± 0.3 and -10.0± 0.3 X I0-5 cm.2fv. sec. Moreover, the oak lignin
possesses a lower mobility than does the white Scots pine lignin. However,
methylation studies with diazomethane have shown that white Scots pine lignin
contains one phenolic group, whereas oak lignin seems to contain three phenolic
27*
Page 868
REPRINT FROM
HANDBUCH DER PFLANZENPHYSIOLOGIE
ENCYCLOPEDIA OF PLANT PHYSIOLOGY
EDITED BY W. RUHLAND
CO EDITORS
E. ASHBY • J. BONNER • M. GEIGER-HUBER • W. 0. JAMES
A. LANG • D. MOLLER • M. G. ST ALFEL T
VOLUME X
SUBEDITORS K. PAECHt AND P. SCHWARZE
SPRINGER-VERLAG/ BERLIN • GOTIINGEN • HEIDELBERG 19,.8
(PRINTED IN GERMANY>
ESPECIAL COMPOUNDS OF LICHENS
BY
S. SIDBATA
Page 869
Handbuch der Pflanzenphysiologie • Encyclopedia of plant physiology
Gesamtdisposition:
Teil I. Allgemeine Grundlagen.
Band I. Genetische Grundlagen physio-
logischer Vorgänge. Konsti-
tution der Pflanzenzelle.
" 11. Allgemeine Physiologie der
Pflanzenzelle.
Teil II. Stoff- und Energiewechsel.
Band 111. Pflanze und Wasser.
" IV. Die mineralische Ernährung der
Pflanze.
"
V. Die 002-Assimilation.
" VI. Aufbau, Speicherung, Mobili-
sierung und Umbildung der
Kohlenhydrate.
" VII. Stoffwechselphysiologie der
Fette und fettähnlicher Stoffe.
" VIII. Der Stickstoff-Umsatz.
" IX. Der Stoffwechsel der schwefel-
und phosphorhaltigen Verbin-
dungen.
"
X. Der Stoffwechsel sekundärer
Pflanzenstoffe.
" XI. Die Heterotrophie.
" XII. DiePflanzenatmung, einschließ-
lich Gärungen und Säurestoff-
wechseL
" XIII. Der Stofftransport in der
Pflanze.
Teil Ill. Wachstum, Entwicklung,
Bewegungen.
Band XIV. Wachstum und Wuchsstoffe.
"
"
XV. Differenzierung und Entwick-
lung.
XVI. Außenfaktoren in Wachstum
und Entwicklung.
" XVII. Physiologie der Bewegungen.
l. Bewegungen durch Einflüsse
mechanischer und elektri-
scher Natur sowie durch
Strahlungen.
2. Bewegungen durch Einflüsse
der Temperatur, Schwer-
kraft, chemischer Faktoren
und aus inneren Ursachen.
" XVIII. Sexualität, Fortpflanzung,
Generationswechsel.
Synopsis of the contents:
Part I. General foundations.
Vol. I. Genetic control of physiological
processes. The constitution of
the plant cell.
" II. General physiology of the plant
cell..
Part II. Metabolism.
Vol. 111. Water relations of plants.
" IV. Mineral nutrition of plants.
" V. The assimilation of carbon
dioxide.
" VI. Formation, storage, mobiliza-
tion, and transformation of
carbohydrates.
" VII. The metabolism of fats and
related compounds.
" VIII. Nitrogen metabolism.
" IX. The metabolism of sulfur- and
phosphorus-containing com-
pounds.
" X. The metabolism of secondary
plant products.
" XI. Heterotrophy and heterotrophic
plants.
" XII. Plant respiration, incl. fermen-
tat.ions and acid metabolism in
plants.
" XIII. Translocation in plants.
Part III. Growth, development,
movements.
Vol. XIV. Growth and growth substances.
" XV. Differentiation and develop-
ment.
" XVI. External factors affecting
growth and development.
" XVII. Physiology of movements.
l. Movements due to mechani-
cal and electrical stimuli
and to radiations.
2. Movements due to the ef-
fects of temperature, gra-
vity, chemical factors and
internal factors.
"XVIII. Sexuality, reproduction, alter-
nation of generations.
Handbuch der Pflanzenphysiologie • Eneyelopedia of plant physiology
Gesamtdisposition:
Teil I. Allgemeine Grundlagen.
Band I. Genetische Grundlagen physio-
logischer Vorgänge. Konsti-
tution der Pflanzenzelle.
" ll. Allgemeine Physiologie der
Pflanzenzelle.
Teil ll. Stoff- und Energiewecbsel.
Band Ill. Pflanze und Wasser.
" IV. Die mineralische Ernährung der
Pflanze.
"
V. Die 002-Assimilation.
" VI. Aufbau, Speicherung, Mobili-
sierung und Umbildung der
Kohlenhydrate.
" VII. Stoffwechselphysiologie der
Fette und fettähnlicher Stoffe.
" VID. Der Stickstoff-Umsatz.
" IX. Der Stoffwechsel der schwefel-
und phosphorhaltigen Verbin-
dungen.
" X. Der Stoffwechsel sekundärer
Pflanzenstoffe.
" XI. Die Heterotrophie.
" Xll. DiePflanzenatmung, einschließ-
lich Gärungen und Säurestoff-
wechsel.
" Xlli. Der Stofftransport in der
Pflanze.
Teil ID. W acbstum, Entwicklung,
Bewegungen.
Band XIV. Wachstum und Wuchsstoffe.
"
"
XV. Differenzierung und Entwick-
lung.
XVI. Außenfaktoren in Wachstum
und Entwicklung.
" XVII. Physiologie der Bewegungen.
1. Bewegungen durch EinflüBBe
mechanischer und elektri-
scher Natur sowie durch
Strahlungen.
2. Bewegungen durch EinflÜBBe
der Temperatur, Schwer-
kraft, chemischer Faktoren
und aus inneren Ursachen.
" XVID. Sexualität, Fortpflanzung,
Generationswechsel.
Synopsis of the eontents:
Part I. General loundations.
Vol. I. Genetic control of physiological
processes. The constitution of
the plant cell.
" ll. General physiology of the plant
cell.
Part ll. Metabolism.
Vol. Ill. Water relations of plants.
" IV. Mineralnutrition of plants.
" V. The assimilation of carbon
dioxide.
" VI. Formation, storage, mobiliza-
tion, and transformation of
carbohydrates.
" Vll. The metaboliBm of fats and
related compounds.
" VIII. Nitrogen metabolism.
" IX. The metabolism of sulfur- and
phosphorus-containing com-
pounds.
" X. The metabolism of secondary
plant products.
" XI. Heterotrophy and heterotrophic
plants.
" Xll. Plant respiration, incl. fermen-
tations and acid metabolism in
plants.
" xm. Translocation in plants.
Part m. Growtb, development,
movements.
Vol. XIV. Growth and growth substances.
" XV. Differentiation and develop-
ment.
" XVI. External factors affecting
growth and development.
" XVII. Physiology of movements.
1. Movements due to mechani-
cal and eleotrical stimuli
and to radiations.
2. Movements due to the ef-
feots of temperature, gra-
vity, ohemical faotors and
internal faotors.
"XVID. Sexuality, reproduotion, alter-
nation of generations.
Page 434
418 F. F. NoRD and G. DE STEVENS: Ligninsand lignification.
C. Infrared absorption spectra.
The infrared spectrum of spruce nativeligninwas first determined by JoNES
(1949 [2]). He found it tobe generally similar to lignins isolated by strong chemi-
cal reagents. Since then the infrared spectra of the native and enzymically
liberated lignins from white Scots pine, oak, birch, maple, bagasse and "kiri"
wood have been determined by NoRD and coworke-rs. In all cases, they were
A
B
c
D
6'?5
Nolive birch Lignin
#olive ookfignin
No live mop/e Lignin
O.~~~~~~~~~LUlU~~UL~LULU~~~~LuLU~uJ
Z.O 5.0 6'.0 7.0 8.0 !l.O 10.0 11.0 1Z.O 1J.O 1'1.0 75.0 16'.0
Wove Lengtll [ Microns]
Fig. 5. Infrared absorption spectra. A White scots pine native Iignin; B birch native Iignin; C oak native Iignin;
D maple native Iignin.
found to resemble closely the spectrum of spruce native lignin. A few of these are
presented in Figure 5.
The band at 3400 cm.-1 is due to the presence of bonded hydroxyl groups in
the lignin complex. However, the intensity of the absorptionband in this region
is less pronounced in the case of bagasse native lignin than in the other lignin pre-
parations. This supports the chemical data in that bagasse native lignin contains
fewer hydroxyl groups than the other lignins. Absorption from 1700 to 1660 cm. -1
is characteristic for the presence of an aldehyde or ketone carbonyl group. This
Page 435
Electrophoresis. 419
band is very weak in the case of maple lignin. Phenyl ring skeletal vibrations with
possible para substitution are evidenced at 1600 and 1510 cm.-1• The 1430 cm.-1
band establishes the presence of aliphatic groupings in the lignin complex. A
difference in relative intensity at 1325 cm.-1 should also be pointed out. Such
variations aremoreevident at 1265, 1255 and 1220 cm.-1, which are in the spectral
region of aromatic or aliphatic unsaturated C-0 groups, and also at 1140, 1163,
1155 and 1125 cm.-1 • These differences possibly are due to different atomic
environments in the lignin complex, and since bands in this region arise from
vibrations of the molecule as a whole, interpretation cannot be specific. The
bands at 2945, 1460, 1380 and 725 cm. -1 are due to Nujol absorption, and thus are
of no interpretative value. Bagasse native lignin and kiri native lignin have sharp
bands at 835 cm. -I, whereas pronounced bands at 890 and 870 cm. -1 are found
in the maple lignin curve. JONES (1949 [3]), using a film technique (evaporation
of a dioxane solution of spruce native lignin) reported bands in the same region
even after prolonged drying. Dioxane is known to absorb in this part of the spec-
trum, but since mineral oil mulls were used with the other lignins, it can safely
be stated that the bands are characteristic of the lignin studied. These bands
are suggestive of strong absorption by trisubstituted phenyl groups. Therefore,
a comparative evaluation of the absorption spectra of these lignins reveals that
they are generally similar, but noticeable differences are observed in the relative
intensities of the bands, suggesting some deviations in the arrangement of groups
present therein. Moreover, it was found that the spectra of the enzymically
liberated lignins from each species of wood were identical with the corresponding
native lignin fractions.
D. Electrophoresis.
The literature is devoid of any studies on the electrophoretic behavior of
lignins. However, due to the presence of at least one phenolic hydroxyl group per
lignin building unit, it appeared that such an investigation would be feasible.
Moreover due to the success in applying electrophoretic methods to the characteri-
zation and fractionation of proteins, a study of the possible applicability of the
method to lignins was undertaken at Fordham.
Preliminary measurements had indicated that the native lignins from white
Scots pine and oak woods are electrophoretically inhomogeneous. Therefore,
the effect of repeated precipitation of lignin on the electrophoretic patterns
was first studied. In Figure 6 are presented the patterns obtained with the oak
lignin after four (1) and eight (2) precipitations. It can easily be recognized
that the oak lignin appears electrophoretically homogeneous, whereas the patterns
obtained with the white Scots pine lignin are characteristic of a multicomponent
system. Whereas occasionally two well defined peaks appeared, the inhomogeneity
revealed itself principally in the existence of trailing boundaries. The inhomo-
geneaus white Scots pine lignin contains components of both higher and lower
mobility than the main components.
The oak lignin was electrophoretically analyzed after every precipitation.
Under identical conditions (PR 10, KCI = 0.1 M, sodium glycinate-HCl = 0.05 M),
the electrophoretic mobility was not affected by the purification to any significant
degree. The mobility of the ascending boundary was found to be always slightly
higher than that of the descending boundary, the respective values being
-10.7 ± 0.3 and -10.0± 0.3 X I0-5 cm.2fv. sec. Moreover, the oak lignin
possesses a lower mobility than does the white Scots pine lignin. However,
methylation studies with diazomethane have shown that white Scots pine lignin
contains one phenolic group, whereas oak lignin seems to contain three phenolic
27*
Page 868
REPRINT FROM
HANDBUCH DER PFLANZENPHYSIOLOGIE
ENCYCLOPEDIA OF PLANT PHYSIOLOGY
EDITED BY W. RUHLAND
CO EDITORS
E. ASHBY • J. BONNER • M. GEIGER-HUBER • W. 0. JAMES
A. LANG • D. MOLLER • M. G. ST ALFEL T
VOLUME X
SUBEDITORS K. PAECHt AND P. SCHWARZE
SPRINGER-VERLAG/ BERLIN • GOTIINGEN • HEIDELBERG 19,.8
(PRINTED IN GERMANY>
ESPECIAL COMPOUNDS OF LICHENS
BY
S. SIDBATA
Page 869
Handbuch der Pflanzenphysiologie • Encyclopedia of plant physiology
Gesamtdisposition:
Teil I. Allgemeine Grundlagen.
Band I. Genetische Grundlagen physio-
logischer Vorgänge. Konsti-
tution der Pflanzenzelle.
" 11. Allgemeine Physiologie der
Pflanzenzelle.
Teil II. Stoff- und Energiewechsel.
Band 111. Pflanze und Wasser.
" IV. Die mineralische Ernährung der
Pflanze.
"
V. Die 002-Assimilation.
" VI. Aufbau, Speicherung, Mobili-
sierung und Umbildung der
Kohlenhydrate.
" VII. Stoffwechselphysiologie der
Fette und fettähnlicher Stoffe.
" VIII. Der Stickstoff-Umsatz.
" IX. Der Stoffwechsel der schwefel-
und phosphorhaltigen Verbin-
dungen.
"
X. Der Stoffwechsel sekundärer
Pflanzenstoffe.
" XI. Die Heterotrophie.
" XII. DiePflanzenatmung, einschließ-
lich Gärungen und Säurestoff-
wechseL
" XIII. Der Stofftransport in der
Pflanze.
Teil Ill. Wachstum, Entwicklung,
Bewegungen.
Band XIV. Wachstum und Wuchsstoffe.
"
"
XV. Differenzierung und Entwick-
lung.
XVI. Außenfaktoren in Wachstum
und Entwicklung.
" XVII. Physiologie der Bewegungen.
l. Bewegungen durch Einflüsse
mechanischer und elektri-
scher Natur sowie durch
Strahlungen.
2. Bewegungen durch Einflüsse
der Temperatur, Schwer-
kraft, chemischer Faktoren
und aus inneren Ursachen.
" XVIII. Sexualität, Fortpflanzung,
Generationswechsel.
Synopsis of the contents:
Part I. General foundations.
Vol. I. Genetic control of physiological
processes. The constitution of
the plant cell.
" II. General physiology of the plant
cell..
Part II. Metabolism.
Vol. 111. Water relations of plants.
" IV. Mineral nutrition of plants.
" V. The assimilation of carbon
dioxide.
" VI. Formation, storage, mobiliza-
tion, and transformation of
carbohydrates.
" VII. The metabolism of fats and
related compounds.
" VIII. Nitrogen metabolism.
" IX. The metabolism of sulfur- and
phosphorus-containing com-
pounds.
" X. The metabolism of secondary
plant products.
" XI. Heterotrophy and heterotrophic
plants.
" XII. Plant respiration, incl. fermen-
tat.ions and acid metabolism in
plants.
" XIII. Translocation in plants.
Part III. Growth, development,
movements.
Vol. XIV. Growth and growth substances.
" XV. Differentiation and develop-
ment.
" XVI. External factors affecting
growth and development.
" XVII. Physiology of movements.
l. Movements due to mechani-
cal and electrical stimuli
and to radiations.
2. Movements due to the ef-
fects of temperature, gra-
vity, chemical factors and
internal factors.
"XVIII. Sexuality, reproduction, alter-
nation of generations.
