Current Biology
Dispatches
in Drosophila. It is clear that Blm plays a
critical role in directing the pathway to
become crossovers. But how, then, is the
activity of Blm (and presumably other
proteins) spatially regulated? It is
tempting — and the worst kind of
sophistry — to suggest that Blm is
regulating the centromere effect and
interference. We have few clues as
to just how these signals might work.
But we now have a molecular target, and
the tools to find its regulators, which
moves us closer to fully understanding
how the centromere effect and
interference work.
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Plant Physiology: Out in the Midday Sun, Plants Keep
Their Cool
Daphne Ezer and Philip A. Wigge*
Sainsbury Laboratory, University of Cambridge, Bateman Street, Cambridge, UK
*Correspondence: Philip.wigge@slcu.cam.ac.uk
http://dx.doi.org/10.1016/j.cub.2016.11.026
Plants use context-dependent information to calibrate growth responses to temperature signals. A new
study shows that plants modulate their sensitivity to temperature depending on whether or not they are in
direct sunlight. This enables them to make adaptive decisions in a complex natural environment.
‘‘It was one of those March days
when the sun shines hot and the
wind blows cold: when it is
summer in the light, and winter in
the shade.’’
— Charles Dickens, Great
Expectations
R28 Current Biology 27, R19–R41, January 9, 2017 ª 2017 Published by Elsevier Ltd.
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Anyone who has hiked in a cool
breeze or spent an afternoon surfing
in sunshine will know that when we’re
kept cool, we sometimes don’t
realise how much UV we’re receiving.
We use heat as a proxy for how
strong the sun is, so when it feels
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the fourth chromosome of diploid females of
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pathway that is partially dispensable in
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14. Kohl, K.P., and Sekelsky, J. (2013). Meiotic
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intermediate metabolism. Mol. Cell 46, 43–53.
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18. Rosu, S., Libuda, D.E., and Villeneuve, A.M.
(2011). Robust crossover assurance and
regulated interhomolog access maintain
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cool we sometimes don’t use as
much sunscreen as we should. A
study in this issue of Current
Biology [1] indicates that plants
do something similar — they
interpret temperature information in
the context of how much sunlight they
Current Biology
Dispatches
are receiving. How and why should
plants adjust their temperature response
based on whether or not they are in the
shade?
Plants are rooted to the spot, and so
must adjust their development to their
environment. Temperature is a key
signal that provides the plant with vital
information about the seasons, and
possible impending stress. Reflecting
this, plant growth and development
are highly responsive to temperature [2].
In particular, warm temperature tends
to increase elongation growth, causing
them to become long and spindly,
as well as to flower early. Previously, it
has been shown that a basic helix-
loop-helix transcription factor,
PHYTOCHROME INTERACTING
FACTOR4 (PIF4), largely controls these
temperature-dependent growth
responses [3,4]. Elongation growth is a
key response of the plant, and many
environmental and endogenous signals
are integrated at the level of PIF4. For
example, the circadian clock regulates
PIF4, and its expression increases in
warm temperatures [5–7]. The activity of
PIF4 protein further integrates
information from gibberellins using
DELLA proteins [8], temperature using
phytochrome B (phyB) [9,10], and blue
light using CRYPTOCHROME 2 (CRY2)
[11]. In this study, the authors
uncover how plants integrate UV-B
information to tune PIF4 activity and
the temperature-dependent growth
response.
Hayes et al. show that the presence
of low levels of UV-B, comparable to
real world levels on a sunny day, are
sufficient to considerably ameliorate
the enhanced activity of PIF4 at 27 C.
Therefore, plants at high temperature
exposed to light containing UV-B have
a strong reduction in their elongation
growth response, as if the plant were
grown in conditions a few degrees cooler.
Moreover, the recently discovered UV-B
photoreceptor UVR8 [12] is required
for this response. UVR8 functions in
a complex with CONSTITUTIVE
PHOTOMORPHOGENESIS1 (COP1)
and SPA proteins, and these factors also
appear to be important for temperature-
dependent growth. It is proposed that UV-
B signalling through UVR8 inhibits both
PIF4 expression as well as the activity of
the PIF4 protein.
This study comes at a burgeoning time
in the field of understanding how light
signals are integrated, particularly in the
context of other environmental signals
such as temperature. Interestingly, a
leitmotif emerging from the studies
is the extent to which whole signal
transduction pathways are considerably
compressed, with the sensor molecule
itself having a major role in the
consequent transcriptomic response
by associating with the promoters of
target genes through interaction with a
transcription factor. This is in contrast to
previous models, where light signals
were proposed to influence gene
expression by activating secondary
messengers within a signalling cascade.
In this case, the activated UVR8–COP1–
SPA complex is proposed to repress the
expression of PIF4. Analogously, it has
recently been shown that a complex of
the active phytochrome B photoreceptor
and likely additional PIF transcription
factors can directly repress the
transcription of growth-promoting
genes such as PIF4 itself and ATHB2.
Similarly, the blue light receptor, CRY2,
associates on the promoters of
responsive genes such as FT [13] in the
presence of activating ligands and can
control gene expression [14] (Figure 1).
Since, for the most part, these light
receptors appear to bind to DNA through
a transcription factor partner, this
arrangement suggests a means to
achieve a considerable degree of
signalling specificity through
programming the expression pattern
and level of the cognate TF. It seems
possible, therefore, that particular
tissues of the plant may indeed perceive
a given light signal differently, depending
on the availability of trans-factors to
communicate the photoreceptor activity
to target genes through promoter
binding.
The concept of context-dependent
signalling is also interesting in terms of
the underlying biology of the response.
Why, for example, does it make sense
for the plant to tune the warm
temperature elongation response to the
level of UV-B? One explanation may be
that plants that experience high UV light
are likely to be in direct sunlight, and
consequently temperatures might be
expected to be high. A warm
temperature signal while in direct
Current Biology 27, R19–R41, January 9, 2017 R29
Re
Bl
UV
d
ue
-B
UVR8 CRY2 PhyB
COP1–SPA PIFs/CIBs PIFs/EC
Flowering Elongation growth
Current Biology
Figure 1. Direct signal transduction of light
information to gene expression in plants.
Plants are able to sense and respond to a wide
range of light wavelengths. In many cases,
photoreceptors perceive light signals and are
recruited to the promoters of target genes through
interactions with cognate transcription factors.
At gene promoters, these signalling complexes
can then directly control the expression of genes
influencing growth and development.
sunlight might be discounted, since this
is the expected condition, so excessive
elongation might not be adaptive. In
contrast, if a plant experiences warm
temperature while in the shade, the
actual temperature in sunlight will be
much higher, and a stronger growth and
developmental response is warranted. In
essence, each temperature
measurement must be interpreted in the
context of the amount of direct sunlight
that portion of the plant receives, in order
to optimise elongation for the
environment.
As well as being of fundamental
value, this study also brings some
helpful practical reminders for lab- and
field-based plant growth. Most plant
growth chambers have little or no UV
light, with most of the light in the
visible part of the spectrum. For some
responses, it might be more realistic and
biologically relevant to include a UV-B
light source to mimic full sunshine. As
well as the lab, crops grown under glass,
which absorbs most UV-B, are noticeably
more susceptible to high temperature
compared with those grown under
plastic. Horticulturalists have been aware
of this phenomenon for many years,
and this result provides the underlying
molecular mechanism. Experimentally,

we often seek to keep conditions as
simple as possible, and minimise
variables. While this helps the
interpretation of results, this study is a
timely reminder that a plant in the field is
growing in a highly complex environment.
Many different variables influence how
plants develop, and their interactions
are often surprisingly complex with
outcomes that are non-linear.
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1. Hayes, S., Sharma, A., Fraser, D.P., Trevisan,
M., Cragg-Barber, C.K., Tavridou, E.,
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(2017). UV-B perceived by the UVR8
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(2009). High temperature-mediated adaptations
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Regeneration: Recorded Live!
Gilbert Weidinger
Institute of Biochemistry and Molecular Biology, Ulm University, Albert-Einstein-Allee 11, 89081 Ulm, Germany
Correspondence: gilbert.weidinger@uni-ulm.de
http://dx.doi.org/10.1016/j.cub.2016.11.022
Salamanders and fish can regenerate amputated limbs/fins. Which cells drive this intriguing process? Two
recent papers have used live imaging of labelled clones of cells to reveal the contribution of connective
tissue.
Appendages regenerate through
formation of a blastema, a population of
proliferative progenitors. To understand
where these cells come from, one
would ideally like to track each
individual cell and their progeny from an
amputated stump into the blastema and
throughout regeneration in live,
unperturbed animals. If this were
combined with reliable ways to
determine the identity of cells, such
comprehensive in toto lineage tracing
could address many intriguing
questions — which cell types contribute
R30 Current Biology 27, R19–R41, January 9, 2017 ª 2017 Elsevier Ltd.
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Ezer, D., Gao, M.J., Khattak, A.K., Box, M.S.,
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to the regenerate, do they regenerate
only their own kind or are some
blastema cells multipotent, is
regeneration driven by stem cells or do
differentiated stump cells contribute? In
addition, questions related to the
behavior of cells in space and time could
be addressed. How is pattern in the
regenerate restored? Hands are different
from lower and upper arms, and this
organization faithfully regenerates. So
where do cells come from that acquire a
certain identity along the proximodistal
(PD) axis? The above envisioned
Current Biology
Dispatches
Phytochromes function as thermosensors in
Arabidopsis. Science 354, 886–889.
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Neme, M., Hiltbrunner, A., Wigge, P.A.,
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(2016). Phytochrome B integrates light and
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Cryptochromes interact directly with PIFs
to control plant growth in limiting blue light.
Cell 164, 233–245.
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definitive in toto live analyses of cellular
behavior during regeneration are not
yet — and maybe never will — be
achievable, but two recent papers by the
Poss and Tanaka groups represent major
steps forward [1,2].
Early lineage tracing studies in
salamanders could not label cells in
unperturbed animals, and rather had to
introduce cells carrying an identifiable
characteristic using tissue grafts, which
might alter cellular behavior during
regeneration. Furthermore, in the
absence of techniques for cell-type