ISSN 10214437, Russian Journal of Plant Physiology, 2010, Vol. 57, No. 3, pp. 305–315. © Pleiades Publishing, Ltd.

, 2010.
Published in Russian in Fiziologiya Rastenii, 2010, Vol. 57, No.3, pp. 323–333.

REVIEWS

Plant Fibers: Initiation, Growth, Model Plants,

and Open Questions1, 2
Simcha LevYadun
Department of Science EducationBiology, Faculty of Science and Science Education,
University of HaifaOranim, Tivon, Israel;
email: levyadun@research.haifa.ac.il
Received July 20, 2009

Abstract—Fibers, which are used as a major raw material in the paper industry, as structural components in
timber for building, and in the manufacture of wooden items, are among the most important renewable
resources. Billions use wood as a major energy source, and fibers are an energyrich component of wood.
They are used for various textiles and as raw material for composites. In this review, I describe the basic char
acters of fibers, their structure, development, uses, and some of the current major model plants for fiber for
mation. I discuss open developmental questions and various aspects of further research. Most of the recent
progress in the biology of fiber formation, especially in their cellwall chemistry, emerged from studies of sev
eral model plants: Arabidopsis thaliana, Populus sp., Eucalyptus sp., flax, and hemp. I stress the critical need
to combine the use of modern methods of research with classical botany. Approaching the issue of fiber for
mation only by molecular or only by classical methods will not only limit the progress, but may result in crit
ical mistakes. Considering the importance of fibers to humanity, it is surprising how little we know about the
biology of fiber formation and how little it is studied as compared, for instance, to the effort to study the
genetics and cell biology of flower organ identity.

Key words: plant  fibers  sclerenchyma  bast  differentiation  wood  intrusive growth  secondary cell wall  hormonal
regulation
DOI: 10.1134/S1021443710030015

1
INTRODUCTION
The term fiber describes several botanical entities:
a mechanical cell type belonging to the sclerenchyma
(true fibers), long cells used to produce paper (true
fibers and tracheids), vascular bundles used for textiles
or to produce ropes (manila and sisal), long trichomes
used for textiles (cotton), and the socalled dietary
fibers. Some of these botanical entities, i.e., scleren
chyma fibers and fibers for paper production, partially
overlap. In this review, I will discuss only “true fibers”,
those belonging to the sclerenchyma.
The sclerenchyma is a simple mechanical plant tis
sue composed of either fibers or sclereids, or both. In
addition, both cell types can be one of the cell types
that build various complicated tissues, such as the
xylem, phloem, and cortex, along with several other
cell types. Alternatively, fibers and sclereids may dif
ferentiate as single cells among many other different
cells, a situation known as ideoblasts [1]. There are
intermediates between fibers and sclereids called fiber
sclereids (e.g., [2]). Fibers serve mostly mechanical sup
1 The article is published in the original.
2 This material was presented at the International
Symposium Plant
Fibers: View of Fundamental Biology, Kazan, Russia, May 28–31,
2009.
port and defense from herbivory, being both hard and flex
ible, but when alive they can also serve in storage [1].
Fibers can be part of both the primary and second
ary plant body. They can be derivatives of the procam
bium (primary phloem fibers, fibers surrounding vas
cular bundles in monocotyledons) [1, 3] or even dif
ferentiate from the ground meristem [4]. In the
secondary body, fibers are mostly cambial derivatives
(fiber bundles in the secondary phloem and fibers of
the secondary xylem) [1, 3]. There are also regenera
tive fibers that differentiate from parenchyma cells
after wounding [5]. Fibers appear in several cellular
types. They can be alive or dead, and uninuclear or
multinucleate [1, 3, 6]. Many fibers have septa that
divide them into several chambers [1, 3]. Fibers can be
short, sometimes less than 1 mm long as in Arabidopsis
thaliana [2], or they may reach a length of 55 cm, as in
Boehmeria nivea (ramie) [7].
There are many patterns of fiber arrangements at
the tissue, organ, and whole plant levels that may sig
nificantly influence the biomechanics of plants, but
this aspect has not been studied much because of the
typical botanist’s limitation in the ability to deal with
theoretical aspects of engineering (but see [8, 9]).

305
306 LEVYADUN
USES OF FIBERS
Fibers are among the most important renewable
resources for humanity. Wood is a major energy source
for many in the world [10]. The energy content of
wood significantly increases when the wood is rich in
highly lignified fibers. At the same time, the mechani
cal properties of timber for construction and as raw
material for various industries are greatly influenced
by their fiber content, chemistry, and structure. The
importance of fibers in the production of textiles is
also well known. From both commercial and biotech
nological points of view, the most important fiber types
are bast (phloem) fibers, mostly used for textiles, and
fibers of the secondary xylem used as the major raw
material for paper production [11]. Their length and
chemistry are critical quality components in the paper
industry [11, 12]. Elimination of lignin from fibers
during paper production is a major cost component in
the paper industry and an important environmental
polluting process [13, 14].
Phloem (bast) fibers are a common source of com
mercial fibers originating in several crops including
Linum usitatissimum (flax), Cannabis saliva (hemp),
Corchorus capsularis (jute), Boehmeria nivea (ramie),
and Hibiscus cannabinus (kenaf) [11, 15–17]. In addi
tion to the traditional uses of fibers mentioned above,
in the last several decades they have been used in the
production of many types of composite materials, but
this broad technological issue is outside the scope of
this essay.
To compensate for the everincreasing demand for
wood fibers for the paper industry and to reduce the
pressure on the dwindling natural forests, more wood
of higher quality will have to be produced on less land
by planting high quality and highly productive trees.
Achieving this goal by traditional studies on tree
genetics, physiology, and development will take more
time than the current global trend of deforestation
allows. The urgent global need to improve tree growth
and quality for the paper industry, to reduce forested
habitat destruction, and to produce biofuel and
sequester carbon make the study of producing more
and better fibers a critical global issue. In this respect,
transgenic technology goes hand in hand with nature
conservation, contrary to the belief of the general pub
lic and some extreme environmental activists.
INITIATION OF FIBERS
Fibers initiate regularly in certain parts of the plant
body, e.g., primary phloem fibers, and this regularity is
useful in studying their development. Classic anatom
ical studies of fiber development were done by Esau
[18], for instance, and reviewed in depth in some of
her books [3, 16]. Parallel studies of the hormonal reg
ulation of fiber formation indicated that various GAs
are involved in determination of fiber length and cell
wall structure [19–21]. A synergistic effect of a combi
RUSSIAN JOURNAL OF PLANT PHYSIOLOGY
nation of auxin (IAA) and GA on secondary xylem
(which includes many fibers) differentiation was
established during the same years [22]. I stress that the
early studies indicating the role of GA in fiber differ
entiation were almost forgotten and are cited very
infrequently. This gives many current scientists the
wrong impression that this level of understanding of
the hormonal regulation of fiber formation was
achieved only in the 1980s.
However, wound effects seemed to influence the
experimental results when wounding was part of the
experiments with hormonal application, and these
effects must be taken into consideration (e.g., [23–25]).
In spite of the wound effects in some of the studies,
these classic studies and others produced the basic
understanding of tissuelevel and celllevel processes
of fiber differentiation, and a general picture of the
hormonal regulation of fiber differentiation emerged.
However, the plethora of genes involved, the delicate
developmental details at the cellular level, and the net
works of regulatory cellular and tissue level signaling of
these complicated processes were impossible to study
in great detail with the classical methods of organismic
biology. Now we have various new techniques that
allow us to address these delicate questions with preci
sion undreamt of not so long ago.
The major unsolved problem concerning fiber ini
tiation is that when we see fiber cells, even at early
stages of differentiation, the cellular factors that deter
mine which cells will acquire a fiber fate may have
already ceased to operate. In some studies there was
confusion between fiber initiation (not demonstrated
in spite of the claims) and fiber lignification, which
was the actual measured factor. Such mistakes should
be avoided. Theoretically, the factors that control later
phases of fiber differentiation (when we can already
identify them as developing fibers) may also operate at
the first stage of fiber cell fate determination, but these
factors have not been identified yet. Identifying these
elusive factors may eventually have a great impact on
the economic aspects of fiber production and also have
important contributions in theoretical aspects of dif
ferentiation.
From the wonderful studies in flower organ identity
regulation or from similar studies on cell fate determi
nation in embryos or in the shoot or root apex, we
know that very delicate signaling networks operate
there. I assume that similar regulatory mechanisms are
involved in the determination of fiber initiation in the
primary phloem, the primary wavy fiber band in inflo
rescence stems of A. thaliana, and in the initiation of
secondary phloem and xylem fibers. The problem is
that because of the importance for industry and trends
in funding, current studies are focused mostly on the
regulation of the chemistry of fiber cell walls and
mechanisms influencing fiber length. For vessel mem
bers, we have the very good Zinnia system that allows
using cell cultures to study many cellular aspects of
differentiation (e.g., [26, 27]). However, we still have
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 PLANT FIBERS: INITIATION, GROWTH, MODEL PLANTS
no such system for fibers (or sclereids). Because there
is no such system, many genomic, proteomic, and
metabolomic studies of fiber development result in
data representing averaged developmental stages and
cell types that were pooled together. Some of these
problems were overcome by extracting fibers of spe
cific developmental stages, but this cannot be done in
all cases.
It is already clear that fiber differentiation is a very
complicated issue. The current studies on fiber devel
opment are usually devoted either to the specific role
of a regulator (hormone or gene), the monomers or
polymers involved in cell wall formation, the genes
that produce the studied components and determine
cell wall structures, compilation of large numbers of
genes involved via genomics, or to identifying the pro
teins involved in the process, indirectly via genomics
or directly via proteomics. There is a great need, only
partly met, to combine these approaches. The lists of
genes and proteins resulting from studies of genomics
or proteomics give us only a crude understanding of
their potential role in fiber differentiation. There is a
need to focus on the cellular and subcellular levels of
actions of each of the many identified genes, their pro
teins, the products of structural proteins, and bio
chemical function of nonstructural proteins. Cur
rently, the methods of RNA or protein harvesting used
in many cases to study fibers are not cell type and
developmental stage specific, resulting in studying a
mixture of cell types and stages of cell differentiation
that may differ significantly in gene expression and
protein function. More specific approaches were used
in initiating cotton “fibers” (e.g., [28]) and should be
used for bast fibers and fibers of the secondary xylem.
Large scale and detailed studies of the relevant tissues
at different developmental stages using in situ hybrid
ization of each relevant gene and immunogold or
immunofluorescence of each of the proteins are
needed. I call this approach cytonomics, similar to
genomics and proteomics. Such tedious but systematic
studies, which may be at least partially mechanized,
may expose potential specific roles of the genes even
before mutant or transgene analyses are done. More
over, some of the genes, which operate at the same
time (concerning specific cell age or developmental
stage) and place, have a good chance of being associ
ated biochemically by the formation of protein com
plexes like many such complexes known in cell biol
ogy. Identifying their associations may shed light on
both their function and mode of operation. Such
largescale studies probably demand an international
consortium similar to those that allowed whole
genome sequencing. However, this may be the best
chance to advance in the understanding of fiber for
mation in large steps.
RUSSIAN JOURNAL OF PLANT PHYSIOLOGY Vol. 57
307
FIBER GROWTH
Fibers usually attain their length in two stages, in
the beginning by symplastic growth and later by intru
sive growth [1, 3, 29, 30]. In certain trees, fibers grow
in length even in the second year after the beginning of
their differentiation [16]. Fibers several millimeter
long are common in many plant species [1, 3, 12, 16],
but fibers in Boehmeria nivea (ramie) may reach a
length of up to 55 cm. These long fibers start as initials
of approximately 20 µm, and over a number of months
grow 27 500 times longer [7]. During this growth, the
fiber tips of ramie, like those of other species, must
pass through the middle lamella between up to tens of
thousands of cells, disrupting a very large number of
plasmodesmata [31–33]. However, there are usually
no wound responses following the intrusive growth of
fibers [34].
Many questions have been raised concerning the
intrusive growth of fibers [34], but there has only been
modest progress in solving them (e.g., [35–37]).
PATTERNS OF FIBER FORMATION
IN SECONDARY TISSUES
The current understanding of the regulation of tis
sue patterning in the secondary xylem and phloem of
plants is inadequate. In many dicotyledons, the axial
system of the secondary xylem is composed of vessel
members, tracheids, fibers, and parenchyma, and the
axial system of the secondary phloem is composed of
sieve elements, companion cells, parenchyma, and
bands of fibers or sclereids (e.g., [1, 38]). Tissue pat
terns in the secondary xylem and phloem have mostly
been studied only from a descriptive point of view
(e.g., [16, 38, 39]) and much less in terms of the regu
lation of pattern formation (e.g., [40]). Patterns of
alternating wood areas rich in vessels and paratracheal
parenchyma with wood areas rich in fibers are com
mon in thousands of woody species (e.g., [38, 41, 42]),
indicating the generality of such phenomena. How
ever, there is considerable variability in the relations
between vessels, parenchyma, and fibers. For instance,
in Acacia albida many vessels are found within broad
bands of parenchyma, but others are found within
fiber bands [43]. The fibersurrounded vessels in Ste
monurus luzoniensis, Rubinia pseudoacacia, Homalium
foetidum, and other examples [41] further indicate the
variability in cellular patterns of fibers, for which we
have no understanding whatsoever of their regulation.
The various patterns of fibers in secondary tissues indi
cate migrating zones of signals that induce either fibers
or other cell types in the cambial region. The fact that
there are zones with and without signals for fiber for
mation is an opportunity to identify the genes involved
in fiber initiation.
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308 LEVYADUN
GELATINOUS FIBERS OF TENSION WOOD
The cell wall of fully differentiated fibers appears in
two gross variations. In the first, normal fibers are sig
nificantly lignified. In the second type, gelatinous
fibers of tension wood (classic reaction wood of dicot
yledons) are not lignified [44]. While the lignin is crit
ical for the mechanical functions of normal fibers in
living plants, when plants are considered as raw mate
rial, the lignin may be either an advantage or burden
depending on intended use. For timber and firewood,
production of lignin adds to the value of the product,
but for the paper industry, lignin must be eliminated
from the product. Therefore, the study of fibers is
greatly influenced by the specific needs of various
funding agencies. The low lignin content of gelatinous
fibers is a natural laboratory for reducing lignin in
fibers, and as such should be fully exploited in research
(e.g., [45]) but has not been studied enough yet.
In general, the current hypothesis is that lower lev
els of polar auxin transport (or auxin level) in the cam
bial region induce differentiation of gelatinous fibers
(e.g., [44, 46]). Gibberellin is also involved in tension
wood formation [47, 48], and probably also ethylene
[49]. In spite of some advances in characterizing the
genes involved in tension wood production (e.g., [50,
51]), the whole issue of the hormonal regulation of
tension wood production and gelatinous fiber differ
entiation is not well understood (e.g., [52]). Since
reduced lignin levels in fibers are very important for
the paper industry, differentiation of tension wood
fibers deserves much more attention than it has
received so far.
USING POLARIZED LIGHT
TO STUDY FIBERS
A very simple method to identify fibers at early (but
not the earliest) stages of differentiation is to examine
the tissues under polarized light. As discussed in Lev
Yadun and Flaishman [4], and in LevYadun et al.
[53], it is common to misidentify nonlignified fibers or
fibers produced among other lignified cell types (e.g.,
[54]). Secondary cell walls usually contain both cellu
lose and lignin. However, identification of special cell
types with secondary cell walls found among other
types in unstained material or by means of common
tissue staining is often difficult, and the use of polar
ized light can significantly help in this task. The
microfibrils of cell walls contain crystalline cellulose,
which is birefringent (it has two refractive indices).
The refractive index to light waves parallel to the long
axis of the cellulose chains and microfibrils is larger
than to light passing at right angles. Light polarized by
passage through a polarizing filter will not pass
through a second polarizer oriented at 90° to the first.
When a piece of birefringent material (in this case a
secondary cell wall) is placed between such crossed
polarizers, so that its axis of major or minor refraction
is at 45° to the planes of polarization of the polarizers,
RUSSIAN JOURNAL OF PLANT PHYSIOLOGY
then the light passing through it becomes polarized
and is able to pass through the second polarizer. The
birefringent material between the polarizers appears
bright to the viewer [55]. I propose routinely using this
simple procedure before a decision is made that a cer
tain tissue produces no fibers (e.g., [53]).
MODEL PLANTS FOR FIBER DEVELOPMENT
General Genetic Comment
The most important model plants for fiber forma
tion are either crop plants (flax and hemp), which
passed a very strong genetic bottleneck and strong
selection of most of their characters, eliminating most
of their variability, or forest trees (Populus and Euca
lyptus) that may in many cases even be clonal. A simi
lar strong selection exists in the common laboratory
genotypes of A. thaliana. In polygenic characters, such
as fiber formation, the action of many of the involved
genes depends on the specific genetic background.
The strong selection that these model plants passed
may result in specific genetic situations, which may
differ in specific molecular, biochemical, and develop
mental characters from other genotypes of these spe
cies or from other taxa.
Fibers in A. thaliana
The emergence of A. thaliana as an established
model plant 20 years ago [56] was a critical moment
for plant biology. This modest, small annual plant
became the plant Drosophila, allowing an explosion of
genetic, physiological, developmental, reproductive,
and evolutionary researches. The completion and
publication of the full sequence of its genome [57] was
probably the single most influential study in plant biol
ogy ever. While the critical role Arabidopsis had in
advancing plant biology in general is common knowl
edge, its role in studying fiber biology was much
smaller.
Secondary wood rich in fibers is essential for
quicker progress in studying the genetic, biochemical,
developmental, and physiological aspects of fiber pro
duction for the paper industry. The use of A. thaliana
for studying fiber development opens enormous
opportunities to study the role of all plant hormones
and their combinations, as well as many other physio
logical and environmental factors (e.g., photoperiod,
temperature, water regime, minerals, vibrations, and
gravity) on fiber formation. Moreover, A. thaliana
allows studying the functions of various genes cloned
from other species, which are much less convenient to
study.
The basics of some wood and fiber production in
A. thaliana were described in Russian before it became a
model plant [58]. This study escaped the attention of the
scientific community both because of the language bar
rier and because of its publication several years before
Arabidopsis became the most important model plant.
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 PLANT FIBERS: INITIATION, GROWTH, MODEL PLANTS
Dolan et al. [59] described the production of a small
amount of secondary wood in the roots of A. thaliana, but
again it caused no shift in the general understanding. The
wrong paradigm was that A. thaliana has no secondary
tissues and so unfortunately it could not be used as a
genetic model for wood and fiber production.
The first intentional attempts to use A. thaliana as a
model for fiber production were made in plants grown
individually in large pots to avoid competition and
selfthinning, and in which the inflorescence stems
were cut every day for two to three months to avoid
monocarpic senescence [2, 60]. This simple method
induced much more secondary wood, fiber, and
sclereid differentiation, especially in roots, but also in
stems, by means of physiological rather than genetic
changes, indicating that A. thaliana can be an excel
lent genetic model for fiber formation [2, 60]. The
basic description of the fiber system in roots and
shoots of A. thaliana is given by LevYadun [2].
A. thaliana has three major fiber systems distinguished
according to their site of differentiation, some of
which have subsystems: (1) bast fibers, produced more
in the main root than in the inflorescence stem or
rosette level stem, a few of which are primary phloem
fibers, while others are short fibersclereids produced
in the secondary phloem of roots and shoots at the
rosette level; (2) long fibers in the secondary xylem of
the main root; and (3) even longer primary and sec
ondary fibers in the wavy band of fibers of the inflores
cence stems [2, 4, 60].
Concerning the wavy ring of lignified fibers in
inflorescence stems, the most studied fibers of
A. thaliana, I give a more detailed description. The
early studies done by LevYadun [2, 60] were con
ducted on mature plants. From these results and from
the confusing captions to plates in various other stud
ies (e.g., [61–63]), it became clear that a developmen
tal study of inflorescence stems, beginning from the
first day of their emergence until they become fully
mature, was needed. Otherwise, as will be discussed
below, there is no way to understand both their struc
ture and various anatomical descriptions found in the
literature. This essential developmental study was
therefore done [4] and indeed illuminated several
complicated developmental aspects of inflorescence
stem anatomy. The inflorescence stem of A. thaliana is
characterized by a wavy ring of lignified fibers inward
to the cortex. In the young stem, primary fibers
develop from the ground meristem, forming the outer
part of the pith as the wavy ring of fibers. In many cases
where A. thaliana is grown on agar or under low light in
growth chambers, only this primary component is
formed. In the mature stem of plants growing under good
growth conditions (individual plants in large pots and
high illumination), the fiber system is comprised of both
these primary fibers and secondary ones that differentiate
around the primary ones, being derivatives of the cam
bium. The cambium formed in the inflorescence stem is
not always continuous. Therefore, the secondary xylem
RUSSIAN JOURNAL OF PLANT PHYSIOLOGY Vol. 57
309
and the secondary parts of the wavy band of fibers may
sometimes form only in sectors at the circumference [4].
However, the progress in understanding the genet
ics and cell biology of fiber production by using
A. thaliana was slower than expected. Some physio
logical studies showed that it is possible to manipulate
fiber formation in A. thaliana by changing growth con
ditions [4] and by external hormonal applications [64].
However, as with many studies of this model plant,
there were expectations for genetic breakthroughs.
One promising mutant that induces alterations in fiber
differentiation is the A. thaliana mutant of the gene
REVOLUTA (REV) [61], which was studied and cloned
in parallel as a different gene INTERFASCICULAR
FIBERLESS1 (IFL1), a name implying that the inter
fascicular fibers do not form [62, 63, 65–68]. These
two putative genes were found to be identical [66]. The
gene encodes a class III homeodomainleucine zipper
protein (HDZIP) and is involved in the regulation of
interfascicular fiber differentiation in inflorescence
stems of A. thaliana [63, 69]. The REV/IFL1 gene was
found to be one of the putative target sequences of
microRNA 165 that cleaves the wildtype REV/IFL1
mRNA [68, 70]. However, the nature of the REV/IFL1
gene turned out to be different than considered for
about a decade. Careful examination of the published
histological figures of the mutant [61–63, 65, 67]
raised the possibility that fibers are actually formed in
the inflorescence stems and that the mutation mainly
alters the secondary cellwall composition rather than
causing fiber absence. To clear this issue, LevYadun
et al. [53] grew plants of the mutant and characterized
the interfascicular fiber band of their inflorescence
stems. They found that the mutant produced the typi
cal primary wavy fiber system of the inflorescence
stems. A simple, but critical aspect in this study was
the examination of the microscopic sections under
polarized light, which clearly demonstrated that fibers
were formed by the mutant. The impression of a total
lack of the wavy band of fibers was found to be a result
of poorly lignified secondary walls in many of the
specimens. This specific gene that reduces lignifica
tion in fibers is of great significance for biotechnolog
ical developments for the paper industry and thus for
the global economy and ecology because of the need to
get rid of the lignin in the process of paper making. An
important conclusion from the studies of the fiber sys
tem of Arabidopsis is that a reliable and detailed know
ledge of the developmental aspects of the studied plant
must be gained before mutants are characterized.
While there was little progress from the use of Ara
bidopsis concerning the genes involved in fiber initia
tion, significant progress was made in the regulation of
the structure and chemistry of the secondary cell wall.
Several transcription factors are involved in the regu
lation of secondary cell walls in the fibers [63, 71–73].
This and progress in understanding other factors
involved in fiber development are essential in order to
control fiber chemistry at the genetic level. This
No. 3 2010
310 LEVYADUN
understanding can allow reducing fiber cost, and
reduce deforestation and environmental pollution by
the paper industry. The irrational public fear of trans
genic plants should also be reduced by education to
allow the progress.
Populus and Eucalyptus
Growing plants of A. thaliana of various genotypes
that produce enough secondary growth for studies on
fiber development need daily clipping of the inflores
cence stems and very strong illumination [60], and it
was expected that a genetic solution would eventually
emerge, as it partly emerged only recently [74]. How
ever, in addition to the use of model herbaceous plants
like A. thaliana that produce some secondary tissues,
real trees are needed to study some of the questions of
fiber formation.
The current progress in making poplar a genetic
model for wood and fiber formation (e.g., [75–81])
and the completion of its genome sequencing [82] are
very important for studying fiber formation, in spite of
the difficulty of genetic analysis in a perennial dioe
cious species. The recent progress in xylem and
phloem transcript profiling in Eucalyptus (e.g., [50,
83–85]) opens further opportunities to understand the
regulation of fiber formation. Both species produce
fibers both in their bark and secondary xylem. More
over, as they are important trees for the paper industry,
there are significant resources for their study. The abil
ity to use Arabidopsis to examine various genes (or
their products) from Populus and Eucalyptus helps to
partly overcome the limitations of trees as model
plants (e.g., [86]). The use of transgenic Populus trees
further supported the understanding of the role of GA
in fiber growth [87, 88].
Flax, Hemp, Rice, Tomato, and Tobacco
Currently, flax seems to be the most studied annual
commercial fiber plant (e.g., [45, 89–93]). Hemp
(e.g., [94, 95]) is also important. Rice, a monocotyle
donous cereal, for which we now have excellent
genetic knowledge, produces large quantities of fibers
in its stems and leaves. Unfortunately, rice is not used
as a classic model plant for fiber production. The crit
ical importance of rice for human nutrition diverts the
research effort from its fibers. However, this can and
should be changed. Since we expect that very soon the
whole genome of several other cereals rich in fibers,
such as sorghum, wheat, and maize, will also be
sequenced, cereals in general may also become good
models for fiber production.
There are no developmental or other biological
reasons not to use other model plant species, such as
tomato and tobacco, for which we have very good
genetic data, to study fiber formation and thus to focus
mostly on plants used for commercial fiber produc
tion. Any model plant that will permit a quicker,
RUSSIAN JOURNAL OF PLANT PHYSIOLOGY
cheaper, and deeper understanding of the biology of
fiber formation should be used. The practical applica
tion of the knowledge to actual crops will be easier
when we have more biological facts.
CLASSIC PHYSIOLOGICAL EXPERIMENTS
SHOULD BE REPEATED
BUT WITH GENOMIC AND PROTEOMIC
APPROACHES
While molecular genetics has enabled unbelievable
progress in understanding plant biology in the last
25 years, even these sharp and powerful tools have lim
itations. When a biological character involves many
genes, it may currently be impossible to use mutants or
transgenic plants to study it. QTL analysis may partly
bridge the gap between monogenic and very compli
cated polygenic characters (e.g., [96]), but even QTL
analysis has its limitations. For instance, the whole
cascade of genes involved in wood and fiber produc
tion in Arabidopsis is not known yet in spite of signifi
cant efforts to study them. Fifteen or twenty years ago,
studying such complicated biological attributes using
mutants in a single gene or similar transgenes was not
practical. Genomics, proteomics, and sequences of
entire genomes were not yet available. Therefore,
using physiology, even if a studied physiological
change was actually a black box, allowed progress in
the study of complicated processes. As already dis
cussed above, attempts were made to use A. thaliana as
a model for wood and fiber production in trees by
using plants, in which the inflorescence stems were
repeatedly cut to avoid monocarpic senescence [60].
This simple method induced much more secondary
wood, and fiber and sclereid differentiation, especially
in the main root, but also in stems, by means of phys
iological rather than genetic changes. It took about
15 additional years to produce a transgenic Arabidopsis
that parallels it in producing secondary tissues [74].
Similar, if not longer delays, in producing the right
genetic combinations are expected for other compli
cated physiological or developmental functions.
The polygenic character of fiber differentiation was
also shown while studying the genetic regulation of
fiber formation in wheat (Triticum aestivum) using the
GAinsensitivity locus Rht1 [97]. One of these poly
genes, a calmodulin gene, was found in a parallel study
conducted for other purposes [98]. The whole cascade
of genes involved in fiber development in wheat is not
yet known.
The ability to fully or partly alter fiber differentia
tion was demonstrated in several systems. In pea
(Pisum sativum), differentiation of primary fibers was
found to be dependent on stimuli originating in young
leaf primordia, but external application of auxin alone
was not a sufficient signal to replace the primordial
and induce fibers [99]. In A. thaliana, submergence
eliminated fiber differentiation in inflorescence stems
[4]. In various tree species, wounding was found to
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 PLANT FIBERS: INITIATION, GROWTH, MODEL PLANTS
arrest fiber production in the secondary xylem for
weeks (e.g., [100]). However, enhanced fiber produc
tion was also shown. It is possible to increase fiber
length (e.g., [19–21, 87, 101] and change its lignin
composition [102] by manipulation of auxin and gib
berellins. All these systems and similar ones, in which
fiber development is altered, open the door to the use
of genomic and proteomic approaches (e.g., [92–94,
103–107]) to identity the cascade of genes involved in
fiber initiation and development, even when critical
mutants and transgenes do not exist.
A good example of the use of classic experiments
are those of Wareing et al. [22], which were recently
repeated in Populus, but with the current ability to
study the transcriptome [78] and give new insights that
were impossible to achieve almost half a century ago.
OPEN QUESTIONS
Almost a decade ago [34], I reviewed various ques
tions concerning fiber differentiation. Here I sum
them up briefly, add new questions, and evaluate the
progress since. (1) What are the signals that induce
fiber initiation? (no progress). (2) Is the determination
of fiber cell fate reversible, and untill what stage? (no
progress). (3) What are the signals that initiate cell
elongation? (no progress). (4) What is the mecha
nism/force enabling intrusive growth? (little progress).
(5) How does the cell tip soften the middle lamella
during intrusive growth? (little progress). (6) Is intru
sive growth the result of tip growth alone, or do other
cell parts take place in the process? (little progress)
(7) How does a cell determine the direction of growth?
(no progress). (8) What brings intrusive growth of
fibers to an end? (no progress). (9) When intrusive
growth occurs, how does the plant sense that the pen
etration into the tissue is not foreign or hazardous
(growth of fungal hyphae or mouth parts of an insect)?
(no progress). (10) What are the mechanisms that
coordinate cellular functions in very long cells such as
fibers? (no progress). (11) What determines fiber cell
wall composition? (moderate progress). (12) Since
many fiber cells are multinuclear, how is their multi
nuclearity regulated? (no progress). (13) Many fibers
have septa, how is septum formation regulated? (no
progress). (14) Many fibers die at the end of differen
tiation, how and why? (little progress).
CONCLUSIONS
Fiber differentiation can be divided into a number
of main stages (determination of cell fate to fiber and
tissue patterning, elongation, deposition of a second
ary, usually lignified cell wall, intrusive growth
between other cells without elicitation of wound
responses, nuclear divisions and formation of coeno
cytes, programmed cell death). All these complicated
but coordinated processes are induced by a combina
tion of hormonal signals, mostly GA and auxin, a fact
RUSSIAN JOURNAL OF PLANT PHYSIOLOGY Vol. 57
311
known for half a century, but with complicated molec
ular interactions that are very far from being under
stood. As can be seen, in spite of a certain progress in
understanding the genes involved in fiber formation
and cell wall structure and chemistry, the general biol
ogy of fiber differentiation is almost as enigmatic as it
was when the basic hormonal regulation of fiber for
mation was revealed decades ago.
An important aspect of studying fiber differentia
tion is the academic programs that produce future sci
entists. We can see a global decline in research and the
study of plant development from the broad organismic
and anatomical points of view. The sharp drop in fund
ing of such studies because they are not always “tech
nically advanced” has reached the point where, with
the retirement of the currently small number of
experts, there will be no new good scientists who
understand the relevant developmental aspects to
replace them. This will certainly damage future
research and progress. This trend should be reversed in
time–now.
ACKNOWLEDGMENTS
I thank the late Tsvi Sachs and Roni Aloni for their
illuminating discussions on the hormonal regulation
of fiber formation, Shahal Abbo for the collaboration
on the genetics and physiology of fiber formation in
wheat, Moshe Flaishman for our long cooperation on
Arabidopsis issues and his important comments on the
manuscript, Sarah Wyatt for the collaboration on Ara
bidopsis, and Ron Sederoff for many discussions on
lignification and forest biotechnology.
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