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Biodeterioration of Wooden Cultural Heritage Organisms and Decay Mechanisms in Aquatic and Terrestrial Ecosystems 1st Edition Anastasia Pournou
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Anastasia Pournou
Biodeterioration
of Wooden
Cultural Heritage
Organisms and Decay Mechanisms in
Aquatic and Terrestrial Ecosystems
Biodeterioration of Wooden Cultural Heritage
Anastasia Pournou
Biodeterioration of Wooden
Cultural Heritage
Organisms and Decay Mechanisms in Aquatic
and Terrestrial Ecosystems
Anastasia Pournou
Dept. of Conservation of Antiquities
and Works of Art
University of West Attica
Athens, Greece
This Springer imprint is published by the registered company Springer Nature Switzerland AG.
The registered company address is: Gewerbestrasse 11, 6330 Cham, Switzerland
To Kalliope Paliatsara and to
George Pournos
Preface
vii
viii Preface
the ecological context of decay, and the general biology of the deteriogenic biota,
while in the following chapters, wood biodeterioration is systematically discussed.
Organisms’ taxonomy, distribution, biology, physiology, niche, and growth condi-
tions, along with their mechanisms, patterns, and diagnostic features of decay, are
examined in detail. The impact of biodeterioration on wood is illustrated and
numerous examples of wooden Cultural Heritage that have suffered biogenic
decay are given, intending to associate the detrimental consequences with the
necessary prevention, preservation, and mitigation strategies.
Effective preservation of wooden artifacts and monuments of historical and
cultural significance requires understanding of the extent of damage as well as the
cause and mechanisms of deterioration. Therefore, this book is hoped to be a useful
handbook for wooden Cultural Heritage professionals and specialists, in order to
assist them when taking preventive and remedial measures, design risk assessment
procedures and disaster planning policies and effectively address the problem of
wood biodeterioration.
By demonstrating wood vulnerability to deteriogens, this book also aims to raise
awareness of the danger and of the need for safeguarding wooden Cultural Heritage
for future generations. Wood can provide a tremendous wealth of information about
past civilizations and cultures and the economic and technological context of
mankind’s history, since it has always been closely linked with mans’ activities.
Finally, although a large amount of information has been examined and about
1500 references have been used, it is expected that in some cases important studies
may have been disregarded and this should be kept in mind. Moreover, it has to be
noticed that the systematics and taxonomy of organisms adopted in this book will
inevitably be soon outdated, considering the continuous discovery of new species
and the rapid progress of molecular phylogenetics.
ix
x Acknowledgments
xi
xii Contents
Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 527
Abbreviations
13 13
C NMR C Nuclear magnetic resonance
13 13
C-TMAH C-labeled tetramethylammonium hydroxide, thermochemolysis
AFM Atomic force microscopy
CCA Copper chrome arsenate
FTIR Fourier transform infrared, spectroscopy
LM Light microscopy
NADH Nicotinamide adenine dinucleotide (reduced form)
NADPH Nicotinamide adenine dinucleotide phosphate (reduced form)
PHB Polyhydroxybutyrate
PLM Polarized light microscopy
RLS Radial longitudinal section
SANS Small angle neutron scattering
SEM Scanning electron microscopy
SFG Sum frequency generation, spectroscopy
TBTO Tributyltin oxide
TEM Transmission electron microscopy
TLS Tangential longitudinal section
TS Transverse section
XRD X-ray diffraction, analysis
xv
Chapter 1
Wood Anatomy, Chemistry and Physical
Properties
Land plants colonized the earth during the Paleozoic era, around 470 million years
ago (Graham 1993; Kenrick and Crane 1997; Wellman et al. 2003). Their ancestors
appears to be green algae (charophyte algae) which evolved to spore-producing and
then to seed-producing plants (Kenrick and Crane 1997; McCourt et al. 2004). Plants
having an organized tissue system consisting of tracheary1 elements are named
vascular plants or tracheophytes. Within this taxonomic superdivision of the plant
kingdom, timber-producing trees are principally found into two Spermatophyta
(seed-bearing plants) divisions (Simpson 2010), the Coniferophyta2 (cone-bearing
plants) (Gernandt et al. 2011; USDA, NRCS 2020) and the Magnoliophyta3
(flowering plants) (Takhtajan 2009; USDA, NRCS 2020) and particularly in the
classes of Pinopsida and Magnoliopsida (dicotyledons), respectively (Fig. 1.1).
Vascular plants, as highly organized organisms, have cells, tissues and vegetative
organs such as the root, the stem and the leaf (Fig. 1.2) (Esau 1977; Cutter 1978).
Wood is a stem tissue and has a multifunctional role consisting in transporting water,
soluble minerals and photosynthesis products, in storing food and in supporting the
stem. For this reason, wood cells’ shape, size, orientation, chemistry and mechanical
strength, have been designed in such a manner to serve effectively this multifaceted
role.
Wood in botany is referred to as secondary xylem,4 because it is formed through
the secondary growth of the plant, during which the stem increases in thickness.
1
Tracheary from trachea (τραχεα) in Greek: duct (vasculum in Latin).
2
Coniferophyta is synonymous to Pinophyta (Gymnospermae clade: naked seed), conifers,
softwoods.
3
Magnoliophyta is synonymous to Anthophyta (Agiospermae clade: seed closed into a fruit),
broadleaves, hardwoods.
4
Xylem from xylon (ξύλoν) in Greek: wood.
Fig. 1.1 The two taxonomic divisions of Coniferophyta and Magnoliophyta, containing timber-
producing trees
Fig. 1.2 Location of leaf, stem and root in a vascular plant. Secondary tissues produced by
cambium during a three-year period. The xylem and phloem cells of the third year (3) are located
next to the cambium, whereas xylem and phloem cells produced during the first year (1) are located
at the inner and outer parts of the trunk, respectively
Inversely, during the primary growth, the plant stem mainly increases in length
producing primary tissues. Secondary xylem is closely associated with the secondary
phloem,5 the second tissue constituting the stem which corresponds to the bark
(Fig. 1.2). Xylem cells are mainly water conducting in contrast to phloem cells
which are primary food conducting (Cutter 1978).
In conifers and woody dicotyledons, both secondary xylem and phloem are
produced by the vascular cambium, which is always found located between the
two (Fig. 1.2). Cambium is an embryonic region (zone) that produces cells by
division, and thus it is a lateral “meristem”,6 (Fahn 1974; Esau 1977; Singh et al.
1987). It originates from the procambium that was formed during the primary growth
of the apical meristem, another meristematic tissue found in the shoot tips.
The vascular cambium during the secondary growth of the plant produces
bilaterally and diametrically, secondary xylem and phloem (Fig. 1.2). The most
recently produced xylem and phloem cells are located next to the cambium, whereas
5
Phloem from phloios (φλoιóς) in Greek: bark.
6
Meristem from meristos (μεριστóς) in Greek: divisible (Singh et al. 1987).
1.2 Xylem Development 3
the oldest ones are found at the inner and outer part of the trunk, respectively
(Fig. 1.2).
The cambial activity in temperate regions occurs during spring and summer,
whereas in tropical climates they may be more than one period of active growth.
Vascular cambium usually consists of two types of primary cells, the fusiform
initials, which are long elongated cells with tapered ends and give rise to the axial
tissue systems of xylem and phloem and the ray initials, which are smaller cuboidal
and almost isodiametric cells that produce the horizontal tissue system of the plant
(Fahn 1974; Esau 1977). When a cambium initial divides, it produces a pair of
identical in morphology cells of phloem or xylem, at a time. One cell of the pair
remains merisomatic, preserving its ability to divide indefinitely, whereas the other
named “mother cell” may also further divide, although not endlessly, i.e. the fusi-
forms usually divide only twice to form four cells (Fahn 1974). The number of
xylem mother cells is always higher than the phloem mother cells and the production
ratio is in average tenfold; however, this varies significantly in species (Fahn 1974;
Esau 1977). Mother cells’ derivatives will then transform and mature to different cell
types which will carry out the various functions in the plant. This developmental
process of cell specialization, during which identical cells become diverse complex
tissues systems, is called “differentiation” (Esau 1977). In woody plants the differ-
entiation of procambial and cambial initials into mature xylem cells is
called xylogenesis (Fukuda 1997; Plomion et al. 2001; Roberts and McCann
2000). The duration of xylogenesis can be as short as 4 days for primary xylem
and from 14 to 21 days for secondary xylem (Myburg and Sederoff 2001).
Xylogenesis of secondary xylem usually includes four stages: (a) cell expansion,
(b) cell wall thickening, (c) lignification and (d) programmed cell death (Fig. 1.3)
(Plomion et al. 2001).
(a) During the first stage, cells will form a primary wall which expands longitu-
dinally or radially so cells will reach their final size and shape and consequently
the various cells types of the wood tissue will be created (Fig. 1.3a) (Demura and
Fukuda 2007). Primary wall at this phase is typically a thin, flexible layer
(0.1–1 μm) consisting mainly of polysaccharides and some structural proteins
(Cosgrove 2005). It forms a mechanically strong network, which though will be
extensible until the end of the cell growth (Cosgrove 1999). This extensibility of
cells appears to be regulated by various proteins such as xyloglucan
endotransglycosylases (XETs), endoglucanases and expansins (Cosgrove
1999, 2005; Cosgrove et al. 2002; Campbell and Braam 1999; Plomion et al.
2001). Cells eventually, via the loosening and expansion of their primary
wall, will acquire their unique functional characteristics and their three-
dimensional orientation inside the wood structure (Plomion et al. 2001).
1.6 Wood Chemistry 15
Table 1.1 The chemical components of wood along with their percentage in wood (based on
Tsoumis 1991; Sjöström 1993; Rowell et al. 2005; Laine 2005)
Polymers 90–99%
Polysaccharides
Cellulose 40–55% Linear polymer of β-D glucose
Hemicelluloses 15–25% Heteropolysaccharides consisting of
(a) Hexoses (i.e. glucose, mannose, galactose)
(b) Pentoses (i.e. xylose, arabinose)
(c) Hexuronic acids (i.e. glucuronic, galacturonic)
Glucans 1% i.e. callose, laricinan, starch
Pectins 1% i.e. galacturan, arabinan, galactans
Lignin 18–35% (a) Guaiacyl lignin (coniferyl alcohol)
(b) Syringyl lignin (sinapyl alcohol)
Oligomers 1–10%
Organic 1–10 (a) Volatile compounds: Terpens
(extractives) (b) Resinous compounds: Resin acids, fatty acids (lipids and
waxes), sterols
(c) Phenolic compounds: Tannins, lignans, stilbenes, flavonoids,
phenols
Inorganic (ash) 0.1–5% Ca, K, Na, Mg oxides
1.6.1 Cellulose
Cellulose was firstly isolated from wood in 1839 by Anselme Payen who also
determined its molecular formula (C6H10O5) and its isomerism with starch (Hon
1994; Klemm et al. 2005). In 1920, Hermann Staudinger discovered that cellulose is
not made up of few small molecules of glucose, as it was believed until then, but it is
a covalently linked high-molecular-weight macromolecule (Hon 1994; Klemm et al.
2005). Cellulose is the main component of the plant cell walls and constitutes
approximately half the dry weight of wood (40–55%) (Tsoumis 1991; Sjöström
1993; Rowell et al. 2005; Stevanovic 2016). It is a semi-crystalline polymer,
insoluble in water and in most solvents, including strong alkali; however, it dissolves
in strong acids (Rowell et al. 2005; Stevanovic 2016).
Cellulose is a long linear homo-polysaccharide, consisting of β-D-glucose9 units,
linked by (1 ! 4) glucosidic bonds (Fig. 1.10). The linkage is formed by the removal
of a water molecule from the hydroxyl groups of two adjusted glucose units, from
the carbon 1 and 4 (Fig. 1.10). Every second glucose unit is rotated approximately
180 and thus the exact repeated unit of cellulose is not glucose, but the disaccharide
cellobiose (Fig. 1.10) (Fengel and Wegener 2003; Klemm et al. 2005).
The chain length of cellulose is expressed by the degree of polymerization (DP),
and plant cellulose has an average DP of at least 9000–10,000 and possibly as high
9
In solution the open-chain form of β-D glucose is mainly present in a cyclic form and is named β-D
glucopyranose.
16 1 Wood Anatomy, Chemistry and Physical Properties
Fig. 1.10 The cellulose macromolecule consisting or repeated units of β-D glucose, linked with 1–4
glucosidic bonds
Fig. 1.11 The supramolecular structure of cellulose, in where cellulose molecules are aggregated
into elementary fibrils, microfibrils and macrofibrils
as 15,000 (Fengel and Wegener 2003; Klemm et al. 2005; Rowell et al. 2005;
Stevanovic 2016). Cellulose molecules have a strong tendency to form intra- and
intermolecular hydrogen bonds between OH-groups (Sjöström 1993; Fengel and
Wegener 2003). Thus, β-D-glucose units are associated together, forming parallel
chains which aggregate and structure isodiametric elementary fibrils with average
diameter between 2 and 4 nm (Fig. 1.11) (Hon 1994; Fengel and Wegener 2003;
Stevanovic 2016). Approximately 40 of these fibrils held together, create a fibrillar
bundle which is called microfibril that has a length of several tens of micrometers
and a diameter ranging from 10 to 25 nm corresponding to more of 200 cellulose
chains (Fig. 1.11) (Hon 1994; Fengel and Wegener 2003; Stevanovic 2016). Finally,
microfibrils will further aggregate to form long fibrils the macrofibrils, which are
the largest structural unit of cellulose with a diameter of about 400–500 nm
(Fig. 1.11) (Fahn 1974; Chen 2014) and can be visible with a light microscope
(Esau 1977).
Microfibrils contained highly ordered and less ordered regions of folded cellulose
chains which are longitudinally arranged and appear to be oriented for native
cellulose (cellulose I) in a parallel direction (Hon 1994; Fengel and Wegener
Another random document with
no related content on Scribd:
“Yes, querida.”
Hand in hand, the lovers left the adobe, and the somber echoing
tunnel, with the electric wires seen like a spider’s web across its
farther end, was to them an underground passage to Paradise.
—Copyright, and used by kind consent of the author.
Note.—Spanish words are pronounced according to the continental
pronunciation, and each vowel is given a syllable. “Si Ma-dre,” pronounced See
Ma´dray, yes, mother. “Ma-ma-ci-ta,” pronounced Ma-ma-cee-tah, little mother.
“Sin Ver-gu-en-za,” pronounced Seen Vehr-goo-ain´tha, shameless. “Que-ri-di-ta,”
pronounced Kay-ree-dee´tah, little love. “Por-ta-les,” pronounced Por-tah´lays,
covered sidewalks. “Gente decente,” pronounced Hen´tay day-then´tay, the
aristocracy. “Coch-i-no,” pronounced Co-chee´no, pig. “Lin-di-ta,” pronounced
Leen-dee´ta, pretty. “Que-ri-da,” pronounced Kay-ree´da, beloved.
Just before daybreak next morning three stealthy figures crept out
and made their way toward Ford’s Creek. One skulked behind the
other two, dogging their steps and taking advantage of the darkness
to keep very near to them. At the grim trysting-place they halted and
were soon joined by other stealthy figures, and together they sat
down to wait for the daylight. The seconds conferred for a few
minutes. The ground was paced off, and a few, low-pitched orders
prepared the young men for business.
“I will count three, gentlemen,” said Lieutenant Custis. “At three,
you are to fire.”
At last daylight came, gray and timid at first, and then red and bold
as the sun came clearly up. The pistols were examined and the men
placed face to face.
“Are you ready, gentlemen?”
But evidently Harrison Randolph was not. He was paying no
attention to the seconds. His eyes were fixed on an object behind his
opponent’s back. His attitude relaxed and his mouth began to twitch.
Then he burst into a peal of laughter.
“Pete,” he roared, “drop that and come out from there!” and away
he went into another convulsion of mirth. The others turned just in
time to see Pete cease his frantic grimaces of secrecy at his master,
and sheepishly lower an ancient fowling-piece which he had had
leveled at Bob Lee.
“What were you going to do with that gun leveled at me?” asked
Lee, his own face twitching.
“I was gwine to fiah jes’ befo’ dey said free. I wa’n’t gwine to kill
you, Mas’ Bob. I was on’y gwine to lame you.”
Another peal of laughter from the whole crowd followed this
condescending statement.
“You unconscionable scoundrel, you! If I was your master, I’d give
you a hundred lashes.”
“Pete,” said his master, “don’t you know that it is dishonorable to
shoot a man from behind? You see you haven’t in you the making of
a gentleman.”
“I do’ know nuffin’ ’bout mekin’ a gent’man, but I does know how to
save one dat’s already made.”
The prime object of the meeting had been entirely forgotten. They
gathered around Pete and examined the weapon.
“Gentlemen,” said Randolph, “we have been saved by a miracle.
This old gun, as well as I can remember and count, has been loaded
for the past twenty-five years, and if Pete had tried to fire it, it would
have torn up all this part of the country.”
Then the eyes of the two combatants met. There was something
irresistibly funny in the whole situation, and they found themselves
roaring again. Then, with one impulse, they shook hands without a
word.
And Pete led the way home, the willing butt of a volume of good-
natured abuse.—From “Folks from Dixie,” copyright by Dodd, Mead
& Company, New York, and used by arrangement.
PART THREE
Melodious Reading
Conversational elements: Pitch, Inflection, Color, Stress, Pause,
Movement, Time. Separate discussions and illustrations with number
of exercises for the pupil to practice. Melody in verse and in prose.
EXPRESSIVE SPEECH[9]
By Robert Lloyd
Exercises in Inflection
By inflection is meant the glide of the voice within a word to a
higher or a lower pitch. This glide may be quick and short, or long
and slow. It may be a rising or a falling glide, or both. The value of
inflection rests in its power to make what is said more emphatic, to
aid in clear enunciation, to aid in overcoming monotony. On all
emphasized words we have an intensified inflection. This is
illustrated in Portia’s speech in “The Merchant of Venice.” In studying
this excerpt we discover that all the emphasized words have a
pronounced inflection. In the first group of words, “If to do were as
easy as to know what were good to do,” we find the most intensified
inflection is upon the word “know” because this is the most emphatic
word of the group. This reveals that inflection is one of the most vital
means of emphasis.
In regard to inflection as an aid to clear enunciation, we find that
inflection occurs upon the accented syllable of a long word, and if
due attention is given to the syllable upon which the accent falls, the
word will receive a more perfect utterance. For instance, we can
readily see in the following words, which are often mispronounced,
the important part that inflection plays in the proper pronunciation of
them:
abdomen
abject
acclimate
address
admirable
alias
brigand
caricature
chastisement
chauffeur
combatant
contumely
demoniacal
discourse
exquisite
finance
grimace
herculean
horizon
impious
impotent
incomparable
indisputable
industry
inexplicable
interpolate
inquiry
lyceum
mausoleum
mischievous
obligatory
research
resource
superfluous
traverse
vagary
vehement
vehicle
virago
verbose
virtue
virtually
Kinds of Inflection
Falling Glide in the voice indicates a complete and positive
assertion. For example:
“The Prince’s banner wavered, staggered backward,
hemmed by foes!”
I find earth not gray but rosy, heaven not grim but fair of hue.
Do I stoop? I pluck a posy. Do I stand and stare? All’s blue.
—Browning.
I must have left my book on this table last night. (Read two ways.)
There are three pleasures pure and lasting, and all derived from
inanimate things—books, pictures, and the face of nature.
—Hazlitt.
What right have you, O passer by the way, to call any flower a
weed? Do you know its merits? Its virtues? Its healing qualities?
Because a thing is common, shall you despise it? If so, you might
despise the sunshine for the same reason.
Oh, yes, I begin to remember you now. Do you really think it true?
Now clear, pure, hard, bright, and one by one, like the hailstones,
Short words fall from his lips fast as the first of a shower,
Now in two-fold column: Spondæ, Iamb, Trochee,
Unbroken, firm-set, advance, retreat, trampling along,—
Now with a sprightlier springiness, bounding in triplicate syllables,
Dance the elastic Dactylics in musical cadences on;
Now their voluminous coil intertangling like huge anacondas,
Roll overwhelmingly onward the sesquipedalian words.
—Browning.
Resolve!
To keep my health!
To do my work!
To live!
To see to it that I grow and gain and give!
Never to look behind me for an hour!
To wait in weakness and to walk in power;
But always fronting onward to the light.
Always and always facing toward the right.
Robbed, starved, defeated, wide astray—
On, with what strength I have!
Back to the way!
A Study of Pitch
Pitch is simply the modulation of the voice as high or low. In
natural speech we seldom have more than one word on the same
pitch. Note the constant change of pitch in a good conversationalist.
In listening to such, we discover what?
First: If one idea is expressed on one pitch, its antithesis is
instinctively expressed on another pitch. For example: “When our
vices leave us, we flatter ourselves we leave them.” “The prodigal
robs his heir, the miser robs himself.” “Excess of ceremony shows
want of breeding.”
Second: A quick leap of the mind causes a leap in the voice, or, in
other words, it causes a change of pitch. For example: “So you say
you are going to—Well, hello, John! How did you get here?”
There can be no definite rules laid down governing Changes of
Pitch. If we think progressively, giving ourselves completely to each
successive idea, permitting our movement of tone to be the direct
outcome of the action of the mind we shall have no difficulty in
modulating our pitch.
In reading the following selections, note carefully the natural
tendency of the voice to change pitch as the mind leaps from one
thought to another.
O larks, sing out to the thrushes,
And thrushes, sing to the sky!
Sing from your nests in the bushes,
And sing wherever you fly.
Study in Stress
If we read or speak aloud naturally and earnestly, there occurs in
our voice a succession of beats or pulsations. If these pulsations
occur at regular intervals, our speech will be “singsong” and
monotonous. Thus:
a
I wandered lonely cloud
as
and
That floats on high o’er hills,
vales
a
When all at once I crowd
saw
o
A host of golden dills.
daff
—Shakespeare.
Abraham Lincoln used scripture quotations very frequently and
powerfully.