COPLBI-857; NO.

OF PAGES 6

Available online at www.sciencedirect.com

Programming desiccation-tolerance: from plants to seeds to

resurrection plants
Jill M Farrant1 and John P Moore2
Desiccation-tolerance (DT) evolved as the key solution to
survival on land by the early algal ancestors of terrestrial plants.
This first DT involved utilizing rapidly mobilisable repair
mechanisms and is still found today in mosses, such as Tortula
ruralis, and ferns, such as Mohria caffrorum. The first seed
plants lost vegetative DT while investing their seeds with
tolerance mechanisms improving their survival in unfavourable
environments. The mechanisms of DT in seeds are strongly
connected to their developmentally regulated maturation
programs. We propose that angiosperm resurrection plants
acquired tolerance by re-activating their innate DT mechanisms
in their vegetative tissues. Here we review the current
hypotheses regarding the genetic evidence for the evolution of
DT in resurrection plants. We also present strong evidence
showing the activation of seed specific genetic elements in the
vegetative tissues of resurrection plants.
Addresses
1
Department of Molecular and Cell Biology, University of Cape Town,
Private Bag, Rondebosch, 7701, South Africa
2
Institute for Wine Biotechnology, Stellenbosch University, Private Bag
X1, Matieland, 7682, South Africa
Corresponding author: Farrant, Jill M (Jill.Farrant@uct.ac.za)

Current Opinion in Plant Biology 2011, 14:16

This review comes from a themed issue on
Physiology and metabolism
Edited by Ute Kramer and Anna Amtmann

1369-5266/$ see front matter

Published by Elsevier Ltd.
DOI 10.1016/j.pbi.2011.03.018

widely evolved the gametophytic solution to desiccation

tolerance and these vegetative tissues are termed fully
desiccation-tolerant as they can withstand very rapid
drying, on the order of seconds to minutes [3]. The main
mechanisms responsible for the DT of mosses are
believed to be related to cellular repair strategies upon
re-hydration. The lack of vascular tissues and sophisticated tissue patterning ensures a rapid and uniform drying
rate upon desiccation in mosses [4]. The molecular processes related to DT in mosses have been most intensively studied in Tortula ruralis (Figure 1A). One of the
key adaptations in Tortula ruralis is the packaging of
desiccation-related mRNA transcripts into mRNPs (messenger ribonucleic protein complexes) with polysomes so
that upon rehydration these transcripts would become
immediately available for the effective repair of desiccation-associated damage [5]. Pteridophytes have been
suggested to show a mixed form of DT, intermediate
between full and modified DT [2]. A recent study has
shown that DT is widespread among fern gametophytes
and related to habitat preference [6]. The simple
morphology of the fern gametophytes governs the
water-holding capacity of the thallus and probably the
DT [6]. Hence it is likely that moss and fern gametophyte
DT is of a similar, if not identical, nature. In the case of
some ferns the sporophyte is stages, as well as the spores,
are desiccation-tolerant, the nature of which is largely
similar to that of the gametophytic tissue. In these tissues
several of the mechanisms of DT reported to occur in
angiosperms (which have modified DT) are present
[7,8,9], confirming an intermediate status of DT. In the
case of the fern Mohria caffrorum (Figure 1C) the DT of
the sporophyte is seasonal, being DT in the dry and
desiccation sensitive (DS) in the wet season [9] and it
would be interesting to know if genetic programming of
the gametophyte is responsible for this switching process.

Introduction the invasion of dry land

Early land plants evolved from aquatic algal ancestors
millions of years ago [1]. One of the first and most
formidable obstacles to the successful adaptation to terrestrial environments is desiccation [1]. Rapid drying
owing to heat, sunlight or wind can cause desiccation
to occur in sensitive vegetative tissues within minutes of
exposure. These desiccation-induced stresses are visible
at all intertidal shore zones where algae are exposed to
rapid changes in water availability throughout a normal
day. Chlorophytic algae, specifically the Choleochetales
group, which gave rise to Charophyceaen species were
the precursors to the Bryophyte and Tracheophyte
lineages of terrestrial plants and must have been the first
groups to acquire desiccation tolerance [2]. Bryophytes
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Physiological and metabolic processes

associated with DT
The changes in hydration levels and cellular stresses
associated with water loss as proposed for seeds [10,11]
are shown in Figure 2. While there has been no research on
water properties in resurrection plants, our research has
shown they experience similar stresses [reviewed in
[12,13]] and the changes in water content and metabolic
responses to desiccation are similar to those proposed for
seeds (Figure 2. [11]). Initial water loss (type V water) is
accompanied by osmotic adjustment to prevent turgor loss.
With loss of type IV water (<0.7 gH2O gDW 1) mechanical stress associated with decreasing cell volume [14,15]
occurs and in seeds this is minimized by progressive
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2 Physiology and metabolism

Figure 1

(a)

(b)

(c)

(d)

cytoplasmic compaction resulting in membrane fusion,

and protein and membrane denaturation is believed to
be prevented by the replacement of water with solutes
capable of substituting for the hydrogen bonds lost owing
to dehydration [2123]. Additional stabilization of the
subcellular milieu is believed to be achieved via cytosolic
vitrification [10,23,24]. Solutes believed responsible for
replacement and stabilization include: 1) sucrose and
oligosaccharides [reviewed in [11,12]] and 2) proteins,
particularly Late Embryogenesis Abundant (LEA)
proteins (reviewed in [11,18,23,25]) and small heat shock
proteins (sHSP) [2528]. Physiological and biochemical
studies on resurrection plants have shown that all of these
changes accompany desiccation in resurrection plants
(Figure 2 [reviewed [12,13]]).

Genetic programs underlying seed DT

(e)

The genetic programs responsible for DT in (orthodox)

seeds must have arisen from the gametophytic and/or
sporophytic tissues of mosses and ferns, the bryophyte
and/or pteridophyte ancestors of seed plants [2]. Not all
seeds are desiccation tolerant and a number of species
produce seeds, termed recalcitrant, that are desiccation
sensitive. Such species occur in environments in which
seeds are released into conditions that are immediately
conducive to germination. It has been proposed that as a
consequence of evolution within such environments
genes for seed DT in such species either have been lost
or are permanently repressed [29]. Comparative studies
between recalcitrant and orthodox seeds are required to
establish the reason for differences in DT.

(f)

(g)

Current Opinion in Plant Biology

Photographs of hydrated resurrection plants. A, Tortula ruralis; B,

Selaginella lepidophylla; C, Mohria caffrorum; D, Xerophyta humilis; E,
Myrothamnus flabellifolia; D, Sporobolus stapfianus; E, Craterostigma
plantagineum. Scale bars are included as an indication of size.

accumulation of storage reserves within vacuoles and

cytoplasm preventing plasmalemma withdrawal and
wall collapse [reviewed in [10,11]]. In resurrection plants,
while complex reserves (protein and lipid bodies and
starch) do not accumulate per se, mechanical stabilization
is alleviated by replacement of water in vacuoles with
compatible solutes and reversible changes to cell wall
architecture [9,12,16,17]. Progressive water loss below
0.45 gH2O gDW 1 (type III water and below) results in
membrane appression and ultimate destabilization, and
increasing metabolic perturbations associated with
free radical production, alcohol and carbonyl emission inter
alia (Figure 2). Damage associated with the latter is minimized by increased activity of house-keeping [18] and
seed specific (e.g. 1-cys peroxiredoxin, see also below)
antioxidants and alcohol and aldehyde dehydogenases
and reductases [18,19,20]. Damage associated with
Current Opinion in Plant Biology 2011, 14:16

A number of transcriptomic and proteomic studies have

been undertaken in order to investigate the genetic
program utilized by orthodox seeds upon acquisition of
DT during their development [19,20,30,31] One of the
most comprehensive transcriptomic studies in seeds has
been performed on Medicago trunculata [19]. Transcript
profiling of mature dry seeds have revealed highly abundant transcripts encoding, among others, LEAs, sHSPs
and peroxiredoxins. Highly expressed functional protein
categories included transcripts involved in signal transduction, development, transcriptional regulation, carbohydrate metabolism, storage protein synthesis, lipid
metabolism, abiotic stress response and photosynthesis.
Proteomics analysis has confirmed that many of the
proteins encoded by these transcripts are highly abundant
in seeds [20,30,31]. In order to differentiate between
genes associated with developmental processes and those
associated with acquisition of DT, osmotic stress was reinduced in radicles of germinating M. truncatula seedlings
by incubation in polyethylene glycol (PEG) and transcripts upregulated during this process were analyzed
[19]. The expression of 16086 M. trunculata genes was
followed of which 1300 genes were differentially
expressed during PEG treatment. These were grouped
into several clusters based on the temporal patterning of
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Programming desiccation-tolerance: from plants to seeds to resurrection plants Farrant and Moore 3

%RWC

Figure 2

AWC g/gDW
MPa

0.45
-3

>0.7
>-1.5

0.25
-11

0.08
-150

-90

-80
Loss of turgor

Mechanical stress

Wilting

Cell Shrinkage

Membrane
appression

Anaerobic
respiration

Protein & membrane structure

destabilized

Unregulated
metabolism

Protein degradation

-70

Damage

-60

Carbonyls emitted

Plasmolysis
Maillard reactions
Free radical
production

-50

ROS production
by autooxidation
Alcohols emitted

-40
Catabolic activity
via enzymes

-30

Membrane demixing
& bilayer transitions

-20

-10

IV

V
Osmotic
adjustment

III

Vacuole filling
Antioxidant production

II

I
plants
seeds &
plants

Response

Sugar accumulation
Water replacement
Accumulation of LEAs & HSPs
Down-requlation of photosynthesis
Leaf folding & pigment accumulation
Wall folding
Current Opinion in Plant Biology

Changes in hydration levels in seeds and resurrection plants with associated physical and metabolic damage reported to occur on water loss (modified
from Vertucci and Farrant, 1995, Berjak et al., 2007 and Farrant et al., 2011). Responses of seeds and plants to these stresses are shown, with those
common to seeds and plants indicated in white and those specific to resurrection plants in grey. The decline in water concentration (g H2O gDW 1) as
a function of Relative Water Content (RWC) for the resurrection plant Xerophyta humilis is shown as a solid line. This trend, however, is typical of
several resurrection plant species studied in our laboratory (e.g. Farrant, 2007) and is similar to the trend for seeds given in Berjak et al., 2007.

the data. Early clustered genes encoded proteins

required for initial protection, including many of the
LEAs and several genes encoding stress and defence
proteins, whereas late clusters consisted of genes
involved in sucrose synthesis, storage protein production
and downregulation of metabolism. The importance of
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sucrose was confirmed as 4 a fold increase occurred with

reestablishment of DT [19]. A functional proteomics
strategy for analyzing M. trunculata seeds was also undertaken on the heat stable proteome extracted [20]. Comparative analysis of desiccation-tolerant versus
sensitive imbibed seeds identified 15 proteins showing
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4 Physiology and metabolism

seed-specific expression, including 6 LEA proteins. We

propose that it was this genetic programming in seeds,
probably consisting mainly of the early-stage genes
involved in protection that were not de-activated during
germination and seedling growth that gave rise to the
numerous lineages of resurrection plants during angiosperm evolution and radiation.

The evolution of resurrection plants

All modified-desiccation tolerant plants are seed plants
and therefore seeds are likely to be the source of genetic
programming for the evolution of all angiosperm resurrection plants [2]. Different desiccation-tolerant resurrection plant lineages exist and therefore, acquisition of
seed DT must have occurred multiple times during
angiosperm evolution. Owing to the lack of a fully
sequenced resurrection plant genome transcriptomic studies have been limited to the analysis of expressed
sequence tags (EST) [17]. Microarray and small-scale
sequence cDNA analysis has been performed on a number of species, including, Tortula ruralis [5], Selaginella
lepidophylla [32], Xerophyta humilis [33], Myrothamnus flabellifolia [34], Sporobolus stapfianus [35] and Craterostigma
plantagineum [36]. Figure 1 shows all these species in the
hydrated state. The most comprehensive transcriptomic
analysis of a resurrection plant has recently been performed for C. plantagineum using pyro-sequencing technology [37]. Four cDNA libraries were constructed from
fully hydrated, 48 h dehydrated, 15 day dehydrated and
24 h rehydrated leaf tissue of C. plantagineum. After
sequence assembly over 15,000 UniProt identities were
obtained, the highest coverage to date for any resurrection
plant. The 500 most variable transcripts, across all experimental samples, were partition clustered and subjected
to functional enrichment analysis. Gene ontology (GO)
categories enriched within the six expression clusters
included responses to abiotic stimuli such as ABA, stress
response pathways, oxidative processes, antioxidant
responses to oxidative metabolism, cellular polysaccharide metabolism and cell wall organization, and photosynthesis and cytoskeletal organization. This study
confirmed the results of many previous molecular studies
of DT in this species and others; re-enforcing the importance of LEAs, sugars, antioxidants and cell wall genes
encoding expansins and xyloglucan endotransglucosylases during desiccation [12,13,38]. One interesting observation from the data is the many chromosome scaffold
genes are responsive to desiccation and one hypothesis is
that C. plantagineum has evolved proteins aiding the
recruitment of transcripts to histone complexes during
desiccation and thus utilizes a similar strategy of mRNP
production to that of Tortula ruralis [37]. However we
know that the DT of T. ruralis is a constitutive repair
strategy that is metabolically expensive. By contrast, DT
in C. plantagineum probably evolved from a developmental-seed program that was re-activated to respond to
environmental cues. A molecular signature of seeds in
Current Opinion in Plant Biology 2011, 14:16

resurrection plants, however, has not yet been convincingly demonstrated [18]. One molecular study, currently
unpublished, has produced data that strongly supports the
seed origin of the DT genetic program in angiosperm
resurrection plants [39]. It was reasoned that one
approach to prove the seed origin was to compare gene
expression of desiccated vegetative tissue with mature
dry seed material sourced from a resurrection plant. A
control parallel experiment involving use of a desiccationsensitive plant in order to compare water stressed vegetative tissue with its corresponding mature seed transcriptome was included. In addition to profiling for potential
new pathways, this approach could identify pathways
common to desiccated vegetative tissue and mature dry
seeds. This experiment was carried out using Xerophyta
humilis, a resurrection plant (Figure 1D), and Arabidopsis
thaliana, the desiccation sensitive control, using cDNA
and Microarray technology [39]. Of the X. humilis genes
analyzed, 46% were found to be differentially expressed
between seed and desiccated vegetative tissue. Cluster
analysis and multivariate techniques revealed that the
transcriptomes of desiccated root, desiccated leaf and
seed tissue were very similar to each other in X. humilis.
This is in contrast to A. thaliana where there is no clear
overlap between gene expression clusters of stressed
vegetative tissue and seed tissue, indicating that the
response to water stress is tissue specific in A. thaliana.
Of particular interest was the identification of a common
set of genes in X. humilis, encoding LEAs, HSPs, peroxiredoxins and storage proteins, that were expressed in
roots, leaves and seeds of desiccated X. humilis, but are
seed-specific in A. thaliana (according to TAIR annotation). The overall conclusions from this study are that
desiccation tolerant angiosperms, such as X. humilis and C.
plantagineum, utilize a seed-specific developmental program that is re-activated in vegetative tissues to protect
against desiccation.

Common patterns of desiccation-tolerance

and outstanding questions for future research
In light of the considerable similarities in mechanisms of
DT in seeds and vegetative tissues of resurrection plants
(reviewed above) it is likely that angiosperms acquired
the initial DT programming from seeds but had to co-opt
or adapt to a whole plant situation by developing
additional specific mechanisms, for example, to deal with
desiccation-associated photosynthetic damage and mechanical stress associated with plasmolysis and cytorhesis
(Figure 2). Understanding the interplay between seed
and angiosperm DT mechanisms will inform our
approach to utilize knowledge obtained from resurrection
plants to improve agricultural crop species. Outstanding
questions still in need of answering include:
1. Is early land plant DT of monophyletic origin?
2. Do all orthodox seeds possess the same types of
genetic DT mechanisms?
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Programming desiccation-tolerance: from plants to seeds to resurrection plants Farrant and Moore 5

3. Do different angiosperm resurrection plant lineages

possess different genetic tolerance mechanisms and
can these be traced to their seed progenitors?
4. How many different DT genetic programs evolved?
5. How are seed genes activated in vegetative tissues?

A good recent review of physiological processes involved in desiccation

tolerance in seeds and resurrection plant species.

Answering these questions, and others, will help to elucidate the scientific mechanisms behind the remarkable
DT abilities of seeds and resurrection plants.

16. van der Willigen C, Mundree SG, Pammenter NW, Farrant JM:
Mechanical stabilisation in desiccated vegetative tissues of
the resurrection grass Eragrostis nindensis: does an alpha TIP
and/or sub-cellular compartmentalization play a role? J Exp
Bot 2004, 55:651-661.

Conflict of interest statement

17. Moore JP, Le NT, Brandt WF, Driouich A, Farrant JM: Towards a

systems-based understanding of plant desiccation tolerance.
Trends Plant Sci 2009, 14:110-117.
A good summary review on the key mechanisms proposed for imparting
desiccation-tolerance in a variety of resurrection plant species.

The authors are not aware of any biases or conflicts of

interest that might be perceived as affecting the objectivity of this review.

Acknowledgements
We thank Keren Cooper and Hanlie Nell for invaluable assistance in
compiling Figure 2.

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(Department of Molecular and Cell Biology, University of Cape Town,
South Africa) and Co-Supervised by Katherine J. Denby (Warwick Life
Sciences, University of Warwick, UK) and Jill M. Farrant (author of this
article).

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