The Plant Cell, Vol. 27: 1565, June 2015, www.plantcell.org ã 2015 American Society of Plant Biologists.
All rights reserved.
IN BRIEF
The Elegant Simplicity of the Liverwort Marchantia polymorpha
Plants have evolved several ways to remain
dormant until the time is right for growth. For
example, seeds maintain dormancy through
the antagonistic effects of the phytohor-
mones abscisic acid (ABA) and gibberellic
acid. Auxin (indole-3-acetic acid [IAA]), in
turn, regulates ABA-induced seed dormancy
through its interaction with the ABA signaling
pathway (Liu et al., 2013). Dormancy is also
crucial for non-seed-bearing plants such as
liverworts, a basal lineage of land plants. In
the liverwort Marchantia polymorpha, tiny,
disc-like propagules (of the haploid gameto-
phyte generation) called gemmae lay dor-
mant in gemma cups on the thallus of the
parent plant until they are dispersed by rain-
fall and released from their dormant state,
a process that has long fascinated botanists.
Early studies employing techniques such as
removing gemmae from their cups, applying
exogenous auxins and antiauxins, and care-
ful microscopic observation have suggested
that gemma dormancy is imposed, at least in
part, by auxin emanating from the thallus
apical meristem of the parent plant (LaRue
and Narayanaswami, 1957). Until recently,
the details of this process, including how the
auxin in the thallus is biosynthesized, have
remained a mystery.
In angiosperms, most auxin is derived
from Trp via indole-3-pyruvic acid (IPyA) in a
two-step pathway catalyzed by numerous
auxin biosynthetic enzymes encoded by
multigene families, with TRYPTOPHAN AMI-
NOTRANSFERASE OF ARABIDOPSIS (TAA)
enzymes catalyzing the Trp-to-IPyA conver-
sion and YUCCA (YUC) enzymes catalyzing
the IPyA-to-IAA conversion. The recent de-
velopment of M. polymorpha as a model
system (including the availability of mutants,
a transformation system, and bioinformatic
tools) has allowed Eklund et al. (2015) to
uncover the elegant simplicity of the auxin
biosynthetic pathway in this species. The
authors found that M. polymorpha contains
only one TAA gene and two YUC genes,
www.plantcell.org/cgi/doi/10.1105/tpc.15.00431
Knockout of the TAA gene. Wild-type plant (left)
and TAA knockout line taa#190 (right) grown for
2 weeks on medium without (upper panels) or with
(lower panels) IAA. Bars ¼ 5 mm. (Reprinted from
Eklund et at. [2015], Figure 4A.)
making this IPyA-dependent auxin biosynthe-
sis toolkit the simplest known in any land plant.
The authors constitutively expressed
YUC1 and YUC2 in M. polymorpha, pro-
ducing plants that appeared similar to
those treated with high levels of exogenous
auxin. Constitutive expression of TAA pro-
duced a similar (but milder) phenotype, with
YUC or TAA transcript levels in the trans-
genic lines positively correlated with the
severity of the phenotype. YUC overex-
pression was correlated with increased IAA
levels, as expected. These findings leave
little doubt that the biochemical roles of
TAA and YUC1/2 in the IPyA pathway are
conserved between liverworts and angio-
sperms. Moreover, the observation that
TAA overexpression had a much milder
effect than YUC overexpression suggests
that conversion from IPyA to IAA by YUC2 is
the rate-limiting step in this pathway during
normal M. polymorpha thallus development.
Expression analysis revealed that YUC1 is
sporophyte specific, while YUC2 and TAA
are expressed in both the sporophyte and
thallus. Experiments employing GUS fused
to the TAA or YUC2 promoter revealed that
these genes are expressed in meristematic
tissue, i.e., shoot apical regions of thalli and
gemma cup bases, the main sites of IAA
production. Dormant gemmae produced
almost no GUS signals, suggesting that
little or no IAA is produced in the gemmae
via the IPyA pathway. The authors then
eliminated or reduced the contribution of
the IPyA pathway to the total IAA pool in
M. polymorpha gametophytes through the
creation of various TAA knockout and
YUC2 knockdown lines, which exhibited
severe developmental defects that were
partially rescued by the application of IAA
(see figure). Strikingly, when gemmae did
form in the gemma cups of these lines, they
did not maintain dormancy. These findings
indicate that auxin is essential for gameto-
phyte development and that it promotes
gemma dormancy. They also demonstrate
the promise of M. polymorpha as a simple,
elegant model system.
Jennifer Lockhart
Science Editor
jlockhart@aspb.org
ORCID ID: 0000-0002-1394-8947
REFERENCES
Eklund, D.M., et al. (2015). Auxin produced by
the indole-3-pyruvic acid pathway regulates
development and gemmae dormancy in the
liverwort Marchantia polymorpha. Plant Cell
27: 1650–1669.
LaRue, C.D., and Narayanaswami, S. (1957).
Auxin inhibition in the liverwort Lunularia. New
Phytol. 56: 61–70.
Liu, X., Zhang, H., Zhao, Y., Feng, Z., Li, Q.,
Yang, H.Q., Luan, S., Li, J., and He, Z.H.
(2013). Auxin controls seed dormancy through
stimulation of abscisic acid signaling by in-
ducing ARF-mediated ABI3 activation in Ara-
bidopsis. Proc. Natl. Acad. Sci. USA 110:
15485–15490.
 The Elegant Simplicity of the Liverwort Marchantia polymorpha
Jennifer Lockhart
Plant Cell 2015;27;1565; originally published online June 2, 2015;
DOI 10.1105/tpc.15.00431
This information is current as of September 2, 2015

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