Hornwort - An Overview - ScienceDirect Topics

12/4/2019 Hornwort - an overview | ScienceDirect Topics
Hornwort
Related terms:
Cyanobacterium, Plant Virus, Symbiont, Nitrogen, Sporophyte, Cycadophyta,

Nostoc, Moss, Liverwort, Potato
Nanoparticle Ecotoxicology
Ashok K. Singh PhD, in Engineered Nanoparticles, 2016
5.3.4.2 Bioaccumulation of Metal Nanoparticles in Aquatic

Organisms
Metal nanoparticles interact with different components of the natural water
ecosystem that may change the nanoparticles' physical or chemical properties (e.g.,
size, surface charge), resulting in modulation of their environmental behavior and
toxicity Kennedy et al., 2008). Zhang et al. (2012a,b,c) have shown that, in the
aquatic system, the zeta potential of ceria nanoparticles changed from 30.9 mV to
near the zero charge, which may weaken the interparticle electrostatic repulsion
and ultimate aggregation. The dissolved nanoparticles are available to the
ecosystem organisms such as hornwort, mud snails, and fish.
Figure 54 shows the accumulation of nanoparticles in fish. The nanoparticle
concentration increased and peaked to about 700 ng/g early, then decreased and
remained stable at around 300 ng/g throughout the experiment. This may account
for various toxic effects of nanoparticles to fish (Handy et al., 2008). Gill, gut, liver,
and brain tissues are identified as the target organs. Ostroumov and Kolesov (2010)
has shown that gold nanoparticles with or without protein modification can
accumulate in hornworts—a submerged, free-floating rootless aquatic macrophyte
in fresh water.
https://www.sciencedirect.com/topics/pharmacology-toxicology-and-pharmaceutical-science/hornwort 1/11
Figure 54. (A) Accumulation of nanoCeO2 in fish (1), hornwort (2), snail (3), and
water (4). (B) The time course of bioaccumulation (as bioaccumulation factor) is
shown in different species.
Reprinted from Zhang et al. (2012a) with permission.
Zhang et al. (2012a) have shown fast absorption and desorption of ceria
nanoparticles in hornworts. The nanoparticle concentration quickly increased,
peaked at about 45,000 ng/g, and then decreased and remained stable at around
1000 ng/g throughout the experiment (Figure 54(A2)). Direct and quick absorption
of nanoparticles from the sediment surface might result in the high
bioaccumulation (Figure 54(B)) in snails. In water, the ceria-nanoparticle
concentration decreased rapidly (Figure 54(A4)). After 15 days, only 0.095% of the
total ceria was found in the water, indicating that almost all of the nanoparticles
have been thoroughly removed from water. Because most of the applied ceria
nanoparticles were recovered from the sediments, the decrease in the nanoparticle
concentrations is due to their aggregation.
Zhang et al. (2012a) used a static freshwater ecosystem, representing lakes and
reservoirs that are usually stratified and experience little mixing. The hydrodynamic
and morphological characteristics of static ecosystem may be different from those
of marine or estuarine systems. Estuarine and marine systems are continuous
flowing systems with changing salinity. Colloids will precipitate when entering into
an estuarine system due to the change of salinity. Therefore, in a realistic aquatic
ecosystem, the behavior and fate of nanoparticles will be more complicated than
the present simulated system.
Cyanobacteria in Nitrogen-Fixing Symbioses

Edder D. Bustos-Díaz, ... Angélica Cibrián-Jaramillo, in Cyanobacteria, 2019
3.2 Facultative Cyanobacteria Symbionts

Cyanobacterial species are capable of forming facultative symbiotic relationships
with many organisms, especially plants. The host recruits facultative symbionts
during the so-called infection stage from its surroundings. Once the cyanobacteria
contacts the host, the symbiosis is established. To infect the host, symbionts must
be capable of detecting and responding to chemical signals produced by the host
(Meeks and Elhai, 2002). The majority of cyanobacterial facultative symbioses
involve filamentous cyanobacteria, which are capable of developing hormogonia, or
motile cells (Meeks et al., 2002). Since many hosts produce, or are thought to
produce, hormogonia-inducing factors (HIFs), the symbiont must have the
necessary genetic equipment to produce the motile cell and detect the HIF
gradient that will guide them to the host to form the symbiosis (Meeks and Elhai,
2002). The genes encoding this function, then, are crucial to separate strictly free-
living symbionts from their symbiotic counterparts. The capacity to alternate
between symbiotic and free-living lifestyles of facultative symbionts is reflected by
an increase in their genome size, caused by the accumulation of the necessary
genes to sustain both lifestyles. Other typical genomic characteristics of these
symbionts are a high percentage of coding genes, as well as AT content (Medina
and Sachs, 2010). Among the facultative symbioses we find the following:
Chaetoceros-Calothrix rhizosoleniae: The exact mechanism of cyanobiont
infection in the symbiosis between the diatom Chaetoceros and the filamentous
cyanobacteria Calothrix rhizosoleniae is not known. However, this Calothrix
species is believed to be an opportunistic symbiont, as suggested by its genetic
traits. In this symbiotic system, the cyanobacteria are attached to the host,
positioned alongside the diatom’s intercellular spaces, anchored to the host
through its sturdier cells, the heterocysts (Villareal, 1992). It stands to reason
that this contact point allows the exchange of fixed nitrogen. However, an exact
description of this process, if true, is still lacking.
The genome of Calothrix rhizosoleniae SC01 has 5.9 Mbp, an AT percentage of
60.5%, and 90.6% of its genes are predicted to be functional (Foster et al.,
2010). C. rhizosoleniae SC01 has certain nif genes, nifH, and nifK, interrupted by
insertion elements of at least 20 kbp (Hilton et al., 2013). While the genome is
relatively small (just marginally bigger than the genome of the obligate
symbiont Trichormus azollae 0708), the rest of its features are consistent with
those of a facultative symbiont. It is interesting to note how different this
symbiosis is, when compared to the previously mentioned diatom-
cyanobacteria symbiosis. While the cyanobiont of Chaetoceros is located outside
the cell, the cyanobionts of both Hemiaulus and Rhizosolenia are located
intracellular. This changed the evolutionary history of these symbionts, as
revealed by their genomes (Hilton et al., 2013).
Bryophytes-Nostoc: Among Bryophytes, the liverworts Blasia and Calvicularia,
along with hornworts Anthoceros, Phaeoceros, Notothylas, and Dendroceros, have
well-defined symbiotic associations with cyanobacteria (Meeks, 2005; Adams
and Duggan, 2008). The best-studied symbiosis of this group is Anthoceros
punctatus-Nostoc. This symbiosis is formed when Nostoc infects the plant,
which is attracted to its host after release of HIFs into the environment. Upon
infection, the cyanobiont locates itself in slime cavities present on the
gametophyte thallus. When Nostoc enters this region, the cavity closes itself to
prevent further infection, creating the low oxygenic conditions necessary for
enhanced nitrogen fixation. When the symbiosis is established, the quantity of
heterocysts rises sharply, up to 45%–50%, nitrogenase activity increases, and
the photosystem shut down. Most of the nitrogen fixed by the cyanobiont
(80%) is transferred to the host (Adams and Duggan, 2008), via an
uncharacterized mechanism.
There are nine genomes of Nostocales species associated with Bryophytes.
Nostoc sp. Moss2, Moss3, and Moss4 were isolated from Pleurozium schreberi;
Nostoc spp. Moss5, and Moss6 were isolated from Hylocomium splendens; and
Nostoc spp. KVJ2, KVJ10, KVJ20, and KVS11 were isolated from Blasia pusilla
(Warshan et al., 2018). The size of these genomes varied considerably, with
ranges from 7.14 to 10.39 Mb, with an average of 8.69 Mb. The AT content of
these genomes, on the contrary, seems more stable, with values in the range of
58% to 59%, with an average of 58.8%. The majority of these species, with the
exceptions of Nostoc spp. Moss3, Moss4, Moss5, and Moss6, were able to form
functional symbioses with Gunnera manicata (Warshan et al., 2018).
Interestingly, the bryophyte Anthoceros punctatus was able to form a functional
symbiosis with a cyanobacterium isolated from a Cycadophyta (Enderlin and
Meeks, 1983). While there are no studies on the structure of the nif operon of
these microorganisms, the smallest genomes of this group, Nostoc spp. Moss5
(7.24 Mb) and Moss6 (7.14 Mb), were phylogenetically grouped (Warshan et al.,
2018), indicating that they may have similar evolutionary symbiotic histories.
Cycadophyta-Nostoc: All of the known species of cycads can form a symbiosis
with cyanobacteria. The symbiosis is physically located at specialized nodules
called coralloid roots, which host the cyanobionts forming a ring called the
“cyanobacterial zone.” (Costa and Lindblad, 2002) The method of infection is
not known, but HIFs, along with phenolic compounds, are thought to play an
important role in the infection and subsequent symbiosis with the cyanobiont
(Lobakova et al., 2004). Once inside the coralloid root, nitrogen fixation
increases, as well as the proportion of heterocysts (Ahern and Staff, 1994). The
increase in the amount of heterocysts is controlled by the host, via chemical
signaling, greatly increasing the yield of fixed nitrogen. The optimum
percentage of heterocyst per filament seems to be around 60%. When this
threshold is passed, the heterocysts separate themselves from the filament,
becoming inactive (Norstog and Nicholls, 1997). Different cyanobacterial
strains can be found in each nodule of the coralloid root, indicating that the
symbiosis is not specific—at least taxonomically (Costa et al., 1999; Zheng et
al., 2002), but it remains to be seen if there is selection toward a functional
cyanobacterial community. It has been shown that nitrogen fixed by the
cyanobiont is exported from this cyanobacterial zone to the meristem, from
where it can be transported to the rest of the plant (Pate et al., 1988).
Nostoc punctiforme ATCC 29133 (PCC 73102) was isolated from the coralloid
roots of a Macrozamia spp. host (Meeks et al., 2001). The genome of this
symbiont is remarkably big (9.06 Mbp), with regular AT percentage (58.6%) and
a high percentage of coding DNA (81.5%) (Moraes et al., 2017). N. punctiforme,
as expected for a facultative symbiont, is capable of living outside its putative
host, since it has a fully functional photosystem, aside from a complete nif
operon (Meeks et al., 2001; Moraes et al., 2017). In this operon, the gene nifD
presents a longer insertion of an excision element than that found in other
heterocystous species (Moraes et al., 2017). Nostoc punctiforme, however, does
not have the excision elements commonly found at the fdxN and hupL genes
(Lindberg et al., 2000). Whether these features are characteristic of facultative
cyanobionts or are unique to this particular organism remains to be seen.
Three more genomes, from strains that remain to be taxonomically
characterized, have been obtained from coralloid roots of Mexican Dioon plants
(Gutierrez-Garcia et al., 2018). These include strains “Cyanobacterium” T09,
“Cyanobacterium” RF31YmG, and “Cyanobacterium” 106c, which are closely
related to Nostoc species. These strains were isolated from Dioon caputoi (T09)
and Dioon edule (106c and RF31YmG), and similar to Nostoc punctiforme ATCC
29133, they have big genomes and medium AT percentages. Other symbiotic
organisms, both closely related to Nostoc species and noncyanobacteria, were
found to be present in the coralloid roots of these cycads (Gutierrez-Garcia et
al., 2018), which is in agreement with a recent report of the Asian cycad Cycas
bifida (Zheng et al., 2018). Finally, there is also a draft genome of a Nostoc
symbiont isolated from a Japanese Cycas revoluta in 1977. The genome of this
isolate, named Nostoc cycadae WK-1, had a length of 6.99 Mb, and a AT
percentage of 59.6 % (Kanesaki et al., 2018).
Gunnera-Nostoc: Similar to cycads, every Gunnera (40 species) form symbiotic
associations with cyanobacteria (Bonnett and Silvester, 1981). This association
is physically located in the stem, near the leaf base, in glands overflown with
mucilage (Silvester and McNamara, 1976). The infection process is well
documented and culminates with the penetration of cortical gland cells of
Gunnera by the cyanobiont (Silvester and McNamara, 1976; Johansson and
Bergman, 1992). Once there, the cyanobiont begins its continuous nitrogen
fixation activity (Silvester and McNamara, 1976). Simultaneously,
photosynthetic activity diminishes, and the symbiont differentiate more
heterocysts per filament, up to 60%–80% (Johansson and Bergman, 1992).
While the cyanobiont is located intracellularly, this relationship is not
transmitted to seedling, which obligates every new generation to form the
symbiosis de novo (Silvester and McNamara, 1976). This facultative symbiosis
is the most effective among all of the symbioses reported to date, with up to
90% of the fixed nitrogen exported to the host, making Gunnera plants totally
impervious to changes in external levels of nitrogen (Silvester et al., 1996).
As with bryophytes, no Nostoc isolated from a Gunnera has been sequenced.
However, it is known that different symbiotic strains exist and that the
symbiosis can be artificially established with Nostoc punctiforme ATCC 29133
(PCC 73102). Indeed, it has been shown that cyanobionts from these plants,
along with cyanobionts from Cycadophyta and Anthoceros, can match the ability
of N. punctiforme ATCC 29133 to form symbiotic relationships with different
hosts (Bonnett and Silvester, 1981).
Plant Development and Evolution

Péter SzövényiManuel WallerAlexander Kirbis, in Current Topics in Developmental
Biology, 2019
5.7 Evolution of meristems

Plant form and architecture are tightly linked to the activity of meristems.
Indeterminate meristems have independently evolved in the gametophyte phase of
bryophytes and ferns, and in the sporophyte of vascular plants. Gametophyte
meristems considerably differ from those of sporophyte meristems, because the
former are composed of a single stem cell, while the latter consist of a single stem
cell, or a group of cells, overlaying several layers of proliferative cells (Ambrose &
Vasco, 2016; Harrison et al., 2007, 2009; Langdale, 2008; Sanders, Darrah, &
Langdale, 2011). Meristematic activity is not recognizable in the fossil record of
early vascular plants. Furthermore, fossils at the divergence of vascular plants and
bryophytes are missing, which makes it impossible to decide whether sporophytic
indeterminant meristems have evolved once or multiple times independently.
Therefore, all information is coming from observations made on extant taxa
Among the extant representatives of the earliest diverged lineages, mosses,
hornworts and some liverworts have multicellular proliferative sporophytic regions,
which may be—to some extent—homologous to proliferative regions of the
indeterminate meristematic regions of extant vascular plants (Bowman, 2013;
Ligrone et al., 2012b; Tomescu et al., 2014). Importantly, hornwort basal meristems
seem to be indeterminate, while moss intercalary meristems show only transient
activity (Bowman, 2013; French & Paolillo, 1975b; Langdale, 2008; Ligrone et al.,
2012a, 2012b; Villarreal & Renzaglia, 2015). It remains to be seen whether these
structures are homologous to one another, and whether their proliferative activity
shares common regulatory activity with that of the proliferative regions of vascular
plant meristems.
The structure of indeterminate meristem activity of sporophyte shoots in
lycophytes and ferns is highly variable, but it usually consists of a single apical cell,
or groups of cells, overlaying a deeper layer of proliferative cells (Ambrose & Vasco,
2016; Frank et al., 2015; Vasco et al., 2016). Expression of key genes of the SAM in
ferns and lycophytes suggests that their sporophytic meristems are multicellular
structures, and their core regulatory mechanisms may be homologous to that of
the flowering plant SAM (Evkaikina et al., 2017; Friedman, 2011). Nevertheless,
transcriptomic evidences rather suggest that vascular plant meristems have
followed divergent evolutionary trajectories, due either to lineage-specific changes
or to independent evolutionary origins (Frank et al., 2015). Nevertheless, how an
apical stem cell (or group of stem cells) evolved in the sporophyte stage is
unknown, but the transient apical activity of moss sporophyte cells may represent
the antecedent of these cells (Bowman, 2013; Tomescu et al., 2014). Transcriptomic
and experimental evidence support the hypothesis that some regulatory
mechanisms of the sporophytic SAM may have been recruited from those present
in the apical cells of gametophytes (Frank et al., 2015; Whitewoods et al., 2018).
Nevertheless, most experimental evidence suggest that this may have been
fragmentary, because deletion of Class I KNOX and WOX transcription factor
genes have no effect on the gametophyte, and Class III HD-Zip (C3HDZ) genes are
not expressed in the gametophyte apical cell of the moss P. patens (Sakakibara et
al., 2008, 2014; Yip, Floyd, Sakakibara, & Bowman, 2016).
Plant Development and Evolution

John W. Chandler, Wolfgang Werr, in Current Topics in Developmental Biology,
2019
3 Apical–basal polarity: A unifying feature with exceptions

A general feature of initial embryonic zygotic divisions is a filamentous, axial
structure or “primitive spindle” (Bower, 1922). The first division plane is mostly
transversal relative to the archegonium neck/venter or the micropyle/chalaza axis in
seed plants and is often symmetrical in lower plant radiations, resulting in two
equally sized daughter cells: a hypobasal cell at the base of the venter and an
epibasal cell toward the archegonium neck. Exoscopic polarity, shown by extant
bryophytes, defines when the embryonic apical pole develops from the epibasal cell
(Fig. 1D); in contrast, endoscopic polarity in most vascular plants is when the
hypobasal cell gives rise to the embryo apical pole (Fig. 1D). An axis inversion
apparently occurred evolutionarily after the basal bryophyte lineage, although some
ferns also show exoscopic polarity. In species of hornwort, i.e., anthocerotophyta,
the first cell division is longitudinal, which indicates that apical embryonic fate can
be established independently of the initial cell division plane (Wardlaw, 1955; Fig.
1D). Endoscopic polarity in angiosperms results from the first transversal or
oblique zygotic division, to give a small apical cell oriented to the chalazal pole of
the megagametophyte or embryo sac, and a large basal cell toward the micropyle
(Wardlaw, 1955).
A general feature of eudicots, exemplified by A. thaliana, is that the chalazal–
micropylar polarity of the egg prepatterns the apical–basal polarity of the zygote
and the prospective primary shoot–root axis. However, in Zea mays (maize) as a
well-characterized monocot, all cell division planes following the first asymmetric
division at the apical pole are random and create a club-shaped transition stage
embryo that consists apically of a small-celled embryo proper, subtended by a
larger-celled suspensor (Chandler, Nardmann, & Werr, 2008), The shoot–root axis
develops obliquely to the apical–basal polarity of the zygote, with the shoot apical
meristem (SAM) at the adaxial face of the embryo proper directing away from the
future endosperm, and the root meristem at the center of the multicellular
boundary with the subtending suspensor.
An additional embryogenesis feature shown by some gymnosperms is a syncytial
phase of several cycles of nuclear duplication without cytokinesis before
cellularization, which results in a cell mass, or tiers of cells, with the upper cells
forming a proembryo and the basal tiers dividing and elongating to form the
suspensor (Cairney & Pullman, 2007). Discrete embryonic and basal suspensor cell
fates at the apical or basal pole, respectively, are thus established after the syncytial
phase (Fig. 1D). Many gymnosperm taxa display either simple polyembryony,
where egg cells within different archegonia are independently fertilized to generate
multiple zygotes within the seed, or cleavage polyembryony, in which proembryos
divide, but only one embryo usually survives to maturity. Cotyledon numbers are
also variable, with Ginkgo biloba and Ephedra trifurca initiating two, but many taxa
have more.
The high diversity of embryogenesis programs within individual taxa and the poor
correlation between embryo type and taxon impede the identification of
morphological embryonic plesiomorphies (ancestral traits). This is exemplified by
debates concerning the orthology of the monocot scutellum and dicot cotyledons,
or of bryophyte rhizoids and higher plant roots. Given the diversity of initial cell
division patterns and deeply branched land plant phylogenies over temporally long
trajectories, the early axial polarity of land plant embryos appears to represent a
common principle and a highly adaptive trait. This raises questions concerning its
common molecular basis, and how evolutionarily informative genetic models such
as A. thaliana are?
Natural Products Structural Diversity-I Secondary

Metabolites: Organization and Biosynthesis
Michel Rohmer, in Comprehensive Natural Products II, 2010
1.13.4.3 Isoprenoid Biosynthesis in Plants: MVA Versus MEP

Pathway: Cross-Talk between the Cytosolic and the Plastidial
Compartments
The allocation of the origin of isoprene units, MVA pathway for the cytosolic
isoprenoids and MEP pathway for the plastidial isoprenoids, is rarely as strict as
described. Evidence for exchanges of prenyl alcohols diphosphates between the two
cellular compartments has been repeatedly reported. Already in the first study on
ginkgo embryos,11 the C20 diterpene skeleton of ginkgolides, derived from the
acyclic geranylgeranyl diphosphate, was shown to be of mixed origin. If the bulk
(∼95%) of the C20 skeleton was effectively derived from MEP pathway, a significant
part was assembled from a C15 moiety (corresponding to farnesyl diphosphate)
with isoprene units derived from MVA pathway completed by a fourth isoprene
unit derived from the MEP pathway. A similar dual origin was observed for the
formation of the diterpene skeletons of phytol with an involvement of the MVA
pathway in the synthesis of the farnesyl diphosphate-derived moiety of phytol,
which is completed by a fourth unit essentially derived from the MEP pathway, in
several liverworts, Heteroscyphus planus,248,249 Lophocolea heterophylla,249 and
Ptychanthus striatus250 and in the hornwort Acanthoceros punctatus251 or for the FPP
moiety of hinokiol, an abietane diterpene, in Torreya nucifera.252 Isoprenoid
skeletons with isoprene units of dual origin have been later reported in many
series.
In C. roseus cell cultures, upon feeding of [1-13C]glucose and careful analysis of the
terpenoid NMR spectra, a significant contribution of the MEP pathway (∼6%) was
found for the formation of the sitosterol skeleton normally derived from the MVA
pathway.253,254 The MEP pathway contribution to the phytosterol biosynthesis was
even of the same magnitude as that of the MEP pathway in Croton sublyratus.255
The isotope distribution upon feeding of 13C-labeled glucose revealed a mixed
origin of the isoprene units in other series. Upon feeding with [U-13C6]glucose, the
phytol of C. roseus cell culture was shown to be derived from both MEP (∼60%)
and MVA pathways (∼40%).254 The C15 sesquiterpene skeleton of bisaboloxide A
and chamazulene from Matricaria recutita is formed from two isoprene units
predominantly derived from the MEP pathway and a third unit of mixed origin,
being derived from both MEP and MVA pathways.256 The sesquiterpene β-
caryophyllene in carrot roots,194 the sesquiterpene blend from vine leaves,257 and
the nerolidyl moiety of 4-nerolidylcatechol from Potomorphe umbellata258 have also
a dual origin as shown from the labeling patterns obtained upon feeding with 2H-
or 13C-labeled MVA, DX, or glucose. In the monoterpene series, a dual origin for
isoprene units has been found for geraniol and (S)-linalool from grape berries259
and (S)-linalool in the leaves of the strawberryFragaria × ananassa.260 In Piper
gaudichaudianum, the hemiterpene dimethylallyl moiety of gaudichaudianic acid,
a p-hydroxybenzoate derivative, is derived from MEP and from MVA pathway as
shown by the incorporation of [1-13C]glucose whereas the geranyl diphosphate-
derived monoterpene chain results from the condensation of DMAPP derived from
both MVA and DMAPP pathways with an IPP unit essentially made through the
MEP pathway.261 Finally, the dolichols, derived from all-cis polyprenols, shown in a
hairy root culture from Coluria geoides were also shown to be of mixed origin after
incorporation of 13C-labeled glucose as a general precursor or 2H-labeled MVA or
DXP as pathway-specific precursors: statistical analysis of the 13C distribution by
mass spectrometry and NMR spectroscopy suggests that in the prenyl chains up to
13 isoprene units are synthesized in the plastids from IPP derived either from MVA
and from MEP pathways and are completed in the cytoplasm with units solely
derived from MVA pathway.262
In tobacco Bright Tellow-2 cell cultures, the two pathways complement one
another.263 Treatment in the low millimolar range with mevinolin, a potent
inhibitor of the key enzyme of the MVA pathway, hydroxymethylglutaryl coenzyme
A reductase, results in growth reduction that is alleviated by the addition of free
DX. Incorporation studies with [1,1,1,4-2H2]DX show that not only plastoquinone
as expected, but also sterols, normally derived from MVA pathway, are both
synthesized through the MEP pathway in the presence of mevinolin. Growth
inhibition by fosmidomycin, inhibiting DXR, the second enzyme of the MEP
pathway, is restored by the addition of MVA to the culture medium;
complementation with [2-13C]MVA proves that in the presence of fosmidomycin
even the prenyl chain of plastoquinone, a representative of the plastid isoprenoids,
is derived from the MVA pathway. Best incorporation yields of the labeled
precursors are obtained by simultaneous treatment with both inhibitors, showing
that at least in this plant system the two pathways can replace each other.
Treatment of A. thaliana seedlings with the same two pathway-specific inhibitors
pointed out the absence of correlation between gene-expression patterns and the
accumulation of the end products of isoprenoid metabolic pathways, suggesting
that posttranscriptional processes may be involved in the regulation of isoprenoid
metabolism264 All these labeling experiments are consistent with the transport
between the cytosol and plastids of C5, C10, and C15 allylic alcohol diphosphates.
This hypothesis is supported by the fact that isolated chloroplasts from three plants
(spinach, Indian mustard, and kale), envelope membrane vesicles, and
proteoliposomes from solubilized proteins of spinach envelope membranes are
capable of efficiently transporting IPP and geranyl diphosphate, and to a lower
extent DMAPP and farnesyl diphosphate265 The weak incorporation of 14C-labeled
MVA that was reported in earlier studies on mono- and diterpene biosynthesis can
now be attributed to metabolite exchanges in the frame of the cross-talk between
the cytosolic and the plastidial plant cell compartments.5
Plasmodesmata and Cell-to-Cell Communication in

Plants
Biao Ding, ... Young-Min Woo, in International Review of Cytology, 1999
A Evolution of Plasmodesmata
The evolutionary origin of higher plant plasmodesmata remains an outstanding
issue. Because green algae Charales are considered to be the most likely ancestors
of land plants (Graham and Kaneko, 1991; Graham et al., 1991; Kenrick and Crane,
1997), an examination of the plasmodesmal structure in this group of algae would
provide a clue to the evolution of plasmodesmata. Existence of plasmodesmata in
Charales is well established. However, the substructural details have not been
resolved clearly. A major issue concerning algal plasmodesmal substructure is
whether an appressed ER is present. Early studies provided conflicting images, in a
large part due to inferior fixation quality of the cell structures (Lucas et al., 1993).
Two groups of workers reinvestigated the issue using improved fixation protocols.
Franceschi et al. (1994) showed that plasmodesmata of Chara corallina are formed
postcytokinetically across an established cell plate. Furthermore, they found no
evidence for the existence of a central structure such as the appressed ER in the
plasmodesmata. Results suggest that the secondary formation of plasmodesmata is
a rather primitive feature and that the appressed ER arose at a later time during
evolution.
Cook et al. (1997) expanded the structural analysis of plasmodesmata to Chara
zeylanica and bryophytesMonoclea gottschei (liverwort), Notothylas orbicularis
(hornwort), and Sphagnum fimbriatum (moss). Liverworts are believed to be basal
to all land plants, with hornworts or mosses as living sister groups to vascular
plants (Kenrick and Crane, 1997). Thus, the approach of Cook et al. (1997) is of
significant value. In contrast to the situation in C. corallina, plasmodesmata in C.
zeylanica appear to be formed at cytokinesis and contain an appressed ER-like
structure. This structure, however, is not a consistent feature. Plasmodesmata in
the three bryophytes invariably contain the appressed ER. Fine variations, however,
exist among the three species. Branched plasmodesmata, perhaps formed via
secondary branching as documented in tobacco (Ding et al., 1992a, 1993; Itaya et
al., 1998), are observed in Monoclea, but not in the other two bryophytes.
Plasmodesmata of Sphagnum have dense staining pattern in the orifice region,
similar to the sphincters observed in a few vascular plants (Olesen, 1979; Badelt et
al., 1994). Cook et al. (1997) suggest that Chara plasmodesmata are less specialized
than the bryophyte counterparts and that complex plasmodesmata evolved in the
ancestor of land plants before extant lineages of bryophytes diverged.
The work of Cook et al. (1997) clearly established that the appressed ER had already
existed in the plasmodesmata of (probably) the earliest land plants. The question
again is whether it had truly evolved in Charales. The differences between the
observations of Franceschi et al. (1994) and Cook et al. (1997) with regard to the
absence or presence of an appressed ER-like structure could be interpreted in a
number of ways. First, such differences might be species dependent. If this were
true, then it becomes very significant that the appressed ER evolved in some Chara
species, but not in others. Second, the differences may be due to the different
fixation protocols used. In general, the high water content of these cells makes it
very difficult to preserve the cell ultrastructure by any fixation procedures. Thus,
either the appressed ER is lost during fixation and subsequent tissue processing
(Franceschi et al., 1994) or the observed appressed ER-like structure represents the
entrapment of ER or even some membrane fragments during fixation (Cook et al.,
1997). This might explain why such a structure is observed in only some, but not
all, plasmodesmata in a given cell wall section. Third, perhaps the appressed ER
had evolved in Chara, but has not become a stable and consistent structure.
Therefore, it will appear in some plasmodesmata but not in others and is also
sensitive to fixation procedures. The labile nature of the appressed ER has been
discussed in Section II,A. Finally, whether the presence of the appressed ER in
plasmodesmata is developmental or cell specific in Chara is not understood,
although there are indications of plasmodesmal modifications during development
(Kwiatkowska and Maszewski, 1986).
To provide further insight into the evolutionary origin of plasmodesmata, fixation
protocols that can preserve algal cell structure well need to be developed to
reinvestigate plasmodesmata in green algal species that have been studied and in
those that have not been studied. For instance, plasmodesmata in Coleochaetales,
another group of green algae considered to be directly related to land plants
(Kenrick and Crane, 1997), deserve a detailed analysis.
It is interesting to note that while Chara species as well as higher plants have
mechanisms to produce secondary plasmodesmata (Franceschi et al., 1994) or to
modify existing plasmodesmata secondarily by branching (Franceschi et al., 1994;
Kwiatkowska and Maszewski, 1986; Ding et al., 1992a, 1993; Itaya et al., 1998),
ferns do not appear to have evolved such mechanisms (Gunning, 1978; Tilney et al.,
1990). Ultrastructurally, however, fern plasmodesmata (Overall et al., 1982; Tilney et
al., 1990) are much more similar to higher plant plasmodesmata by indisputably
having the appressed ER (Ding et al., 1992b; Botha et al., 1993). So have ferns lost
the mechanism to form secondary plasmodesmata or have ferns evolved
plasmodesmata independently? Equally puzzling is the finding that branching of
plasmodesmata, as observed frequently in higher plants and Chara species, is a
feature of plasmodesmata in liverworts, but not of plasmodesmata in the more
advanced bryophytes hornworts and mosses (Cook et al., 1997). Finally, a few
fungal species have plasmodesmata that are ultrastructurally similar to higher plant
plasmodesmata (Lucas et al., 1993). Resolving these puzzles is not only of value to
understanding the evolution pattern of plasmodesmata, but should also be of value
to understanding the detailed aspects of the evolution of plants.
In addition to structural analysis, comparative studies of the biochemical
compositions of plasmodesmata in green algae, bryophytes, and higher plants are
indispensable. A significant step has been taken by Blackman and Overall (1998) to
characterize the biochemical compositions of Chara plasmodesmata. They have
shown, by immunolabeling, the presence of actin and myosin in Chara
plasmodesmata. This mirrors the detection by immunolabeling of actin (White et
al., 1994) and myosin (Radford and White, 1998) in fern and higher plant
plasmodesmata, respectively. If actin and myosin are confirmed by further studies
to be bona fide components of Chara, fern, and higher plant plasmodesmata, how
they function in plasmodesmal transport requires careful studies. Ding et al. (1996)
showed that treatment of tobacco mesophyll cells injected with cytochalasin D and
profilin results in an increase in the plasmodesmal SEL to at least 20 kDa, whereas
Ding and Tazawa (1989) demonstrated that treatment of Chara cells with
cytochalasin E has no effect on intercellular transport. Isolation of protein
compositions and the corresponding genes of plasmodesmata from green algae,
bryophytes, ferns, and higher plants should complement structural approaches in
painting an evolutionary picture of plasmodesmata.
Comparative functional studies on plasmodesmata in various groups of plants are
also required to complement the structural and biochemical approaches. These
include characterization of molecules that can be transported through
plasmodesmata, when, during evolution, plasmodesmata became competent for
such transport and the mechanisms of transport. In this regard, it is significant to
note that TMV MP interacts with and modifies the functions of the intercellular
communication system in cyanobacteria. These photosynthetic prokaryotes grow as
filamentous and multicellular organisms. Microplasmodesmata of approximately
8–20 nm in diameter play an important role in cell-to-cell communication that
regulates cell differentiation and growth (Wolk, 1996). These microplasmodesmata
are probably composed of mainly proteins (Giddings and Staehelin, 1978,1981).
Expression of the TMV MP in Anabaena sp. strain PCC 7120 leads to cell wall
accumulation of the MP and perturbation of cell differentiation and growth
(Zahalak et al., 1995). Further analysis revealed that the MP induces the formation
of filamentous bundles of approximately 150 nm that traverse the septa (Heinlein
et al., 1998b). It is possible that individual filaments within a bundle traverse a
group of microplasmodesmata. Alternatively, a bundle may traverse a septal pore
that is derived from modification of a microplasmodesma or formed de novo in the
presence of MP (Heinlein et al., 1998b). These data suggest that some components
of the intercellular communication system may be conserved in organisms ranging
from photosynthetic prokaryotes to higher plants.
Copyright © 2019 Elsevier B.V. or its licensors or contributors.

ScienceDirect ® is a registered trademark of Elsevier B.V.

Hornwort - An Overview - ScienceDirect Topics

Uploaded by

Document Information

Original Title

Copyright

Available Formats

Share this document

Share or Embed Document

Sharing Options

Did you find this document useful?

Is this content inappropriate?

Copyright:

Available Formats

Hornwort - An Overview - ScienceDirect Topics

Uploaded by

Copyright:

Available Formats

12/4/2019 Hornwort - an overview | ScienceDirect Topics

Cyanobacterium, Plant Virus, Symbiont, Nitrogen, Sporophyte, Cycadophyta,

5.3.4.2 Bioaccumulation of Metal Nanoparticles in Aquatic

Cyanobacteria in Nitrogen-Fixing Symbioses

3.2 Facultative Cyanobacteria Symbionts

Plant Development and Evolution

5.7 Evolution of meristems

Plant Development and Evolution

3 Apical–basal polarity: A unifying feature with exceptions

Natural Products Structural Diversity-I Secondary

1.13.4.3 Isoprenoid Biosynthesis in Plants: MVA Versus MEP

Plasmodesmata and Cell-to-Cell Communication in

Copyright © 2019 Elsevier B.V. or its licensors or contributors.

You might also like