On the Inside

Author(s): Peter V. Minorsky

Source: Plant Physiology, Vol. 154, No. 2, 25,000th Article Commemorative Issue (October
2010), pp. 602-603
Published by: American Society of Plant Biologists (ASPB)
Stable URL: http://www.jstor.org/stable/20779816
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On the Inside
How Low Can Lignin Go?
Reducing the lignin content of plants

has long been a goal of plant biotech

nology. Lignin is an unwanted product

in the pulp and paper industry because

it increases the costs of paper produc

tion and associated water treatments

New Insights into

the Cuticle
The cuticle, the hydrophobic barrier

that covers the aerial surfaces of the

plant, is mainly composed of cutin, a

polyester matrix, and a complex mix
ture of soluble waxes. The primary func
tion of the cuticle is to retard water loss,

necessary for environmental protection.

Moreover, reducing the lignin content of

but it also acts a barrier against patho

more efficient hydrolysis of Polysaccha

rides in biomass and thus facilitate the

gen invasion. In this second capacity,

the cuticle acts as a passive physical

the raw biomass material may allow

production of biofuel. Thus, much re

search has been aimed at the production

of genetically modified trees with al

tered lignin biosynthesis. A slew of
articles in this issue, including an over

view by Li and Chappie (pp. 449-452),

examine some of the negative physio
logical consequences of lowered lignin
content in trees (particularly in trees
with 40% to 50% reductions in lignin
content) and call into question just how
low lignin contents can go before yield is
affected. Kitin et al. (pp. 887-898) report

that the transgenic poplar genotypes

(Populus tremula X Populus alba) with the

lowest lignin content due to antisense

4-coumarate:CoA ligase (4CL) had re

duced xylem specific conductivity com
pared with that of control trees. Very
low-lignin trees contained areas of non
conductive, brown-colored xylem with
patches of collapsed cells and patches of
noncollapsed cells filled with phenolics.
Thus, the reduced transport efficiency
of the transgenic low-lignin xylem was
largely caused by blockages from tylo

ses and phenolic deposits within ves

sels, rather than xylem collapse. Voelker

et al. (pp. 874-886) also found physio

logical abnormalities in field-grown,

barrier to infection and as a poten

tial source of signaling and elicitor
molecules. Two articles in this issue
provide new insights into the role of
plant cuticles in fighting pathogen in
vasions, while a third describes a new
technique for their visualization and
study.
Fungal pathogens that penetrate the
plant tissue directly through the outer
surface, rather than via natural plant
openings or wounds, must pass through
the plant cuticle and epidermal cell wall.
Penetration of the host surface happens
either by appressoria or by enzymatic

hydrolysis. Necrotrophic plant patho

gens such as Botrytis cinerea typically use

the latter strategy. Curvers et al. (pp.

847-860) report that a mutant of tomato

(Solanum lycopersicum) with reduced

abscisic acid (ABA) production (sitiens)

exhibits increased resistance to the

necrotrophic fungus B. cinerea. This re

sistance is correlated with an H202

regulated cell wall fortification response

in epidermis cells that is absent in

tomato with normal ABA production.

Evidence is presented that is consistent

with the idea that ABA deficiency in

the mutant leads to increased cuticle

low-lignin poplars. They conclude that

permeability and altered cell wall

reductions in lignin content do not

increase wood processability such as

endogenous plant ABA levels affect the

they would benefit the production of

liquid biofuels by fermentation. Finally,

Novaes et al. (pp. 555-561) review evi

dence that the balance between lignin

biosynthesis and biomass enhance

ment is mediated, at least in part, at

the level of transcription, and propose
a model to explain the inverse relation

ship between wood formation and lig

nin content in trees.

www.plantphysiol.org/cgi/doi/
10.1104/ .110.900332

composition. These results show that

composition of the tomato cuticle and

cell wall and demonstrate the impor
tance of cuticle and cell wall chemistry
in shaping the outcome of this plant
fungus interaction.
In reexamining the question of the
role of gibberellic acid in the acquisi
tion of systemic acquired resistance

(SAR), Xia et al. (pp. 833-846) have

uncovered evidence for a role for the

cuticle in producing SAR. SAR involves
the generation of a mobile signal in the
primary leaves that, upon translocation

to the distal tissues, activates defense

responses resulting in broad-spectrum

resistance. Previous studies had indi

cated a role for plastidial glycerolipid

biosynthesis in SAR. Specifically, mu
tations in FATTY ACID DESATURASE7
(FAD7), which lead to reduced trienoic

fatty acid levels and compromised

plastidial lipid biosynthesis, have

been associated with defective SAR.

This study shows that the defective

SAR in fad7-l plants is not associated
with a mutation in FAD7 but rather
a second-site mutation in GLABRA1
(GLI), a gene best known for its role in
trichome formation. These results un

derscore a need to reevaluate pheno

types previously studied in the gli

background. The compromised SAR

in gli plants and two other so-called
"trichome mutants" is associated with
impairment in their cuticles, suggesting

an overlap in the biochemical path

ways leading to cuticle and trichome
development. The application of gib

berellic acid not only enhanced SAR in

wild-type plants but also restored SAR
in gli plants. This enhancement in SAR

was associated with increased levels of

cuticular components but not trichome
formation on gli plants, thus implicat
ing the cuticle as an important compo

nent of SAR.
As the two featured articles above
demonstrate, the cuticle is a target ripe

for further study. Thus, the research

community will no doubt be grateful for
a new technique by Weissflog et al. (pp.

604-610) that allows for the in vivo

chemical imaging of epicuticular waxes.

These authors present evidence that

coherent anti-Stokes Raman scattering
microscopy is a highly effective tool for
investigating the epicuticular wax layer

of plants. The chemical selectivity of

anti-Stokes Raman scattering micros
copy opens the doorway for the online
monitoring of chemical and structural
changes in wax structures.

Nucleotide-Induced
Calcium Oscillations
It is now widely accepted that extra
cellular ATP is an important signaling
compound in plants, where it regulates
such processes as root-hair growth,
tropic curvatures, and responses to
abiotic and biotic stimuli. In mamma

602 Plant Physiology?, October 2010, Vol. 154, pp. 602-603, www.plantphysiol.org ? 2010 American Society of Plant Biologists

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All use subject to http://about.jstor.org/terms

lian cells, where extracellular ATP has

also been documented to be an im

require ATP release via exocytotic se

observed in cold-stored tubers from

wild potato germplasm stocks that are

Cold-Induced Sweetening of

gene expression were observed in CIS

resistant potato germplasm. Thus, CIS
and its associated acrylamide problem

portant messenger, two types of pu

rinoceptors, P2X and P2Y, bind

extracellular ATP. The binding of ex

tracellular ATP to P2X receptors opens

Ca2+ channels, whereas the activation

of P2Y receptors stimulates recruit
ment of heterotrimeric G proteins to

resistant to CIS. Only low levels of VInv

Potato Tubers

Potato (Solanum tuberosum) tubers re

quire cold storage to prevent sprouting,

trigger cytoplasmic signaling and gene

minimize disease losses, and supply

Tanaka et al. (pp. 705-719) have

consumers and the processing industry

with high-quality tubers throughout the
year. Unfortunately, the accumulation of

expression. To elucidate the function

of extracellular nucleotides in plants,

examined the effects of several nu

cleotide analogs and pharmacologi

cal agents on cytosolic free calcium
([Ca2+]cyt) changes in transgenic
Arabidopsis (Arabidopsis thaliana)
plants expressing aequorin or the
Ca2+ sensor Yellow Cam?l?on 3.6.
Extracellular ATP induces a rise in

reducing sugars in cold-stored potato

tubers, a phenomenon referred to as
cold-induced sweetening (CIS), is a per
sistent, costly problem for the potato
processing industry. During potato chip

and French fry production, reducing

sugars (primarily Glc and Fru) in the
tubers react with free amino acids in

the level of [Ca2+]cyt in plant cells. A a nonenzymatic Maillard reaction to

mutant lacking the heterotrimeric G produce brown- to black-pigmented
protein G/3-subunit exhibited a re products that are not acceptable to
consumers. Such products also have
markably higher [Ca2+]cyt elevation
in response to all tested nucleotides
elevated amounts of acrylamide, a
in comparison with the wild type.
neurotoxin and potential carcinogen.

These findings are indicative of a

role for heterotrimeric G proteins in

modulating the cellular effects of

extracellular nucleotides in plants.

Developing methods to reduce acrylam

ide in fried potato products has become

an urgent requirement for the potato

distinct time- and dose-dependent

kinetics compared with the purine
nucleotides ATP and GTP. Moreover,

breeding. For example, in contrast to S.

tuberosum, potato chips processed from
tubers of Solanum raphanifolium acces

[Ca^lcyt elevations were distinct be

The addition of ATP to aequorin

potato clones. Genetic engineering also

[Ca2+]cyt oscillations in the presence of

an apyrase inhibitor. However, in the

absence of this inhibitor, such [Ca2+]cyt

oscillations could be localized to specific
root cell layers using Yellow Cam?l?on

accessions are crossed with diploid

offers approaches for reducing CIS of

potato tubers. Bhaskar et al. (pp. 939
948) demonstrate that silencing the

potato vacuolar acid invertase gene

VInv prevents reducing sugar accumu

lation in cold-stored tubers. Potato chips

3.6 fluorescence. Finally, the authors

processed from VInv silencing lines

present evidence that a vesicle traffick

ing inhibitor reduces the oscillations,

and were light in color even when

suggesting that the oscillations may

Toward Perfecting the

"Golden Potato"

Vitamin A deficiency is a widespread

public health problem in developing

countries, where it is a major cause of

childhood blindness and increases sus

ceptibility to infections. /3-Carotene

exhibits the highest provitamin activity,

and there have been several successful

efforts to improve the provitamin A

content in major crops. Previously, the

research group of Diretto et al. (pp.

899-912) has engineered transgenic
"Golden" potato tubers overexpressing

three bacterial genes for /3-carotene

synthesis {CrtB, Cr?i, and CrtY, encod

ing, respectively, phytoene synthase,

of the various components of the

minipathway (transgenes as well as

sions stored for three months at 2?C

expressing seedlings induced distinct

targeted breeding.

been achieved by traditional plant

showed acceptable color. This extreme

cold-chipping phenotype is heritable
when cold-chipping S. raphanifolium

tween CTP and the purine nucleotides.

VInv gene through biotechnology or

phytoene desaturase, and lycopene

the inhibitory effects of antagonists of

mammalian P2 receptors and Ca2+ in

flux inhibitors on nucleotide-induced

may be controlled by suppression of the

processing industry. Some progress to

Exogenously applied CTP caused ward reducing CIS of potato tuber has

elevations in [Ca ]cyt that displayed

low levels of VInv gene expression were

cretion.

showed a 15-fold acrylamide reduction

tubers were stored at 4?C. Comparable,

/3-cyclase). To define the specific role

promoters controlling them) in modu

lating carotenoid biosynthesis in the
tuber, the authors have now examined

the systematic molecular and biochem

ical characterization of transgenic

plants harboring six different combina

tions of the CrtB, CrtI, and CrtY genes
under the control of constitutive (35S)
and tuber-specific Patatin promoters.
As a result of these studies, the authors

have identified several genes, specifi

cally the cytochrome P450 genes (LUTI
and LUT5), that are likely to play a key
regulatory role in "Golden" tubers and
therefore are excellent candidates for

genetic manipulations aimed at the

further optimizing tuber carotenoid
content.

Peter V. Minorsky
Division of Health Professions and Natural Sciences

Mercy College

Dobbs Ferry, New York 10522

603

Plant Physiol. Vol. 154, 2010

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