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NEWS AND VIEWS
of several mineral elements in many of the
mutants identified).
Second, the results from numerous
expressed sequence tag (EST) and genome
sequencing projects demonstrate that most
plant transporters are encoded by multigene
families, the members of which exhibit over-
lapping, but nonidentical, expression pat-
terns. Seldom is it obvious which members (if
any) of a family are crucial, or are at least
major contributors, to the homeostasis of a
specific ion without knowledge of the mineral
element profiles of the appropriate mutants.
Third, many inorganic nutrients are trans-
ported by more than one transporter family
and many plant transporter families trans-
port more than one nutrient. For instance, in
Arabidopsis, K
+
uptake can be mediated by
members of the KUP/HAK/KT transporter
family, as well as by members of the KAT and
AKT channel families, if not also by other
uncharacterized transporters such as those
belonging to the KEA family
6
. Likewise,
although the CAX and NHX transporter fam-
ilies were originally defined on the basis of the
capacity of their prototypes for Ca
2+
/H
+
and
Na
+
/H
+
antiport, respectively, it is becoming
increasingly clear that this is not what all
members of these families do.
Fourth, there are some plant transporter
families (e.g., the MATE family) or subfami-
lies (e.g., the ATH ABC transporter subfam-
ily) whose transport substrates are completely
unknown. These cannot be second-guessed
using homology arguments but are nonethe-
less implicated in ionic homeostasis by the
results from systematic cDNA microarry
analyses
6
.
Fifth, as if this is not enough in the way of
deliverables for the plant community, it
should be appreciated that ionic profiling will
provide strong clues not only to the func-
tional capabilities of transporters or at least to
[A
U: OK?] the functional networks in which
they participate, but also to the components
or upstream or downstream processes that
modulate the expression, activity, or the
membranes (compartments) to which the
transporters [A
U: OK?] localize to exert their
effects. Indeed, this next phase of research in
ionomics should clarify which genes play key
roles in ion regulation, providing a wealth of
biological information, which in turn should
open the door to an exciting array of possibil-
ities for improving human nutrition and
engineering plants for phytoremediation.
[A
U: concluding sentence ok?]
1. Daar, A.S. et al. Nat. Genet. 32, 229–232 (2002).
2. Lahner, B. et al. Nat. Biotechnol. 21, XX–XX (2003).
3. Arabidopsis Genome Initiative. Nature 408, 796–815
(2000).
4. Jarvis, K.E., Gray, A.L. & Houk, R.S. Handbook of
Inductively Coupled Plasma Mass Spectrometry
(Blackie, London, 1992).
5. Raamsdonk, L.M. et al. Nat. Biotechnol. 19, 45–50
(2001).
6. Maathuis, F.J.M. et al. Plant J. XX, XX–XX (2003).
[AU: pr
ovide vol. & page #s]
sucrose or xylose.
A microbial fuel cell is, in essence, a system
that harvests electrons produced during
microbial metabolism and channels them for
electric current generation. Three different
microbial fuel cell designs (types A, B and C)
have been implemented, with varying degrees
of success. In type A fuel cells, artificial redox
mediators capable of penetrating bacterial
cells are added to the culture solution within
the anodic fuel cell compartment, enabling
electrons produced during fermentation or
other metabolic processes to be shuttled to
the anode
3
.Type B fuel cells incorporate
metal-reducing bacteria (e.g., members of the
Geobacteraceae and Shewanellaceae) that
exhibit special membrane-bound
cytochromes capable of transferring electrons
directly to the electrodes (rather than dis-
solved or solid iron(
III) or manganese(IV/III)
species)
4
.And type C fuel cells oxidize fer-
mentation products, such as hydrogen and
methanol, on electrocatalytic electrodes (that
is, chemically modified electrodes capable of
efficiently oxidizing such metabolites)
5
; these
latter reactions are often inhibited by the elec-
trode materials platinum, gold or carbon
themselves.
All three designs have their share of advan-
tages and disadvantages. Type A fuel cells
show rather good current densities but suffer
from a need to use synthetic and often toxic
redox mediators that have to be added to the
cell culture and are nonrecoverable. Type B
fuel cells can work in natural environments
(e.g., sediments); however, the growth rate of
the bacteria and thus the current densities
generated are very low. And although type C
fuel cells produce the highest current densi-
ties and work with simple and easily available
microorganisms (e.g., Escherichia coli), elec-
trocatalytic electrodes require rather expen-
sive surface modifiers that make
implementation harder
6
.
Chaudhury and Lovley have now found in
Rhodoferax ferrireducens a microorganism
that is, compared with Geobacter or
Shewanella species, superior in its glucose-
catabolizing efficiency (Fig. 1). Bearing in
mind that the organism itself needs some of
the glucose for its own biomass production,
an 80% electron yield is indeed a very surpris-
ing and impressive result. In addition, elec-
trons generated by R. ferrireducens are easily
transferred to the anode without the assis-
tance of electron-shuttling mediators, and the
cells grow at a steady rate, which guarantees a
steady supply of electrons and therefore a
consistent current density.
Whereas microbial fuel cells have a long to
go before they compete with more established
NATURE BIOTECHNOLOGY VOLUME 21 NUMBER 9 SEPTEMBER 2003 3
Energy generation and waste disposal are two
key challenges in the quest for sustainable
societies. Electrochemical fuel cells may, in
principle, provide an elegant solution by link-
ing both tasks. On page XX of this issue,
Chaudhury and Lovley
1
show that Rhodoferax
ferrireducens,a metal-reducing bacterium
recently isolated from subsurface sediments
in Oyster Bay, Virginia, USA
2
,provides a con-
stant flow of electrons to simple graphite elec-
trodes in a fuel cell while oxidizing glucose or
other simple sugars. This is a notable achieve-
ment for several reasons: first, glucose has
never before been shown to be microbially
oxidized in a fuel cell at 80% electron effi-
ciency; second, unlike most other het-
erotrophic microbial systems, the culture in
the anodic fuel cell compartment requires no
dissolved electron-shuttling mediators; and
third, the bacteria can generate a steady elec-
tron flow over extended periods of time while
growing on a variety of simple substrates,
such as lactate, acetate, glucose, fructose,
Bacterial batteries
Fritz Scholz & Uwe Schröder
Long-lived fuel cells lacking diffusional electron mediators have been
created that convert simple and abundant sugars into electricity with >80%
efficiency.
Fritz Scholz and Uwe Schröder are at the
Institute of Chemistry and Biochemistry,
University of Greifswald,17489 Greifswald,
Germany.
e-mail: fscholz@uni-greifswald.de
NEWS AND VIEWS
physico-chemical electrical batteries, several
factors provide added impetus to research into
microbial electricity production. First, bacte-
ria are omnipresent in the environment and
are thus adapted to feeding on virtually all
available carbon sources, which makes them
into potential catalysts for electron flow gener-
ation in biofuel cells—extending the range of
possible substrates for electricity generation
from low molecular fuels (e.g.,hydrogen or
methanol) used in conventional fuel cells to
carbohydrates or more complex organic mat-
ter present in sewage, sludge
7
or even marine
sediments
8
.A further argument relates to the
increasing cost of precious metals. These con-
stitute the catalytic core of conventional fuel
cells, and thus one may envision that in the
future biocatalysts like bacteria may become a
serious cost-reducing alternative. Finally, it is
worth noting that bacteria can be described as
self-reproducing, self-renewing catalysts, and
in an ideal case where a simple initial inocula-
tion of a suitable strain could be cultured con-
tinuously in a fuel cell, long-term fuel cell
operation could be initiated and maintained.
This also represents the major advantage of
microbial-based fuel cells over the closely
related immobilized enzyme–based fuel cell
systems in which catalysis happens on surface-
immobilized enzymes of microbial origin. In
these systems, the enzymatic activity dimin-
ishes quickly over time.
Taking advantage of [A
U: OK?] using sim-
ple monosaccharides from waste or other
detritus as substrates for electricity generation
is an important achievement, which leaves one
wondering about possible applications of this
system. As an example, biotechnological
hydrolysis of the di-, oligo-, and polysaccha-
rides in molasses could be effectively coupled
to a R. ferrireducens–driven fuel cell, thus cre-
ating a power plant virtually limitless with
respect to fuel resources. It is very hard to
guess now what the economy of such systems
would be, but the development of efficient
microbial fuel cells, such as the one described
here, is the first step towards this goal.
1. Chaudhury, S.K. & Lovley, D.R. Nat. Biotechnol. 21,
XX–XX (2003).
2. Finneran, K.T., Johnsen, C.V. & Lovley, D.R. Int. J.
Syst. Evol. Microbiol. 53, 669–673 (2003).
3. Park, D.H. & Zeikus, J.G. Appl. Environ. Microbiol.
66, 1292–1297 (2000).
4. Kim, H.J. et al. Enz. Microbial Technol. 30, 145–152
(2002).
5. Karube, I., Matsunaga, T., Tsuru, S. & Suzuki, S.
Biotechnol. Bioeng. 19, 1727–1733 (1977).
6. Schröder, U., Nieβen, J. & Scholz, F. Angew. Chemie
Int. Edn. 42, 2880–2883 (2003).
7. Kim, B.H., Chang, I.S., Hyun M., Kim H.J. & Park
D.H. WO Patent 01/04061 A1 (2001).
8. Bond, D.R., Holmes, D.E., Tender, L.M. & Lovley, D.R.
Science 295 483–485 (2002).
4 VOLUME 21 NUMBER 9 SEPTEMBER 2003 NATURE BIOTECHNOLOGY
Anode
Cathode
C
6
H
12
O
6
(glucose) + 6 H
2
O
12 H
2
O
6 CO
2
+ 24 H
+
+ 24 e
–
Ion exchange
membrane
e
–
e
–
e
–
e
–
e
–
e
–
e
–
e
–
H
+
H
+
H
+
H
+
H
+
H
+
6 CO
2
+ 24 H
+
+ 24 e
–
Rhodoferax
ferrireducens
Figure 1 Rhodoferax ferrireducens microbial fuel cell. R. ferrireducens burns carbohydrates to CO
2
in
the anodic compartment, a process that produces free electrons which are directly captured by the
anode. From there, the electrons are channeled to the cathode, where they reduce oxygen to water. The
transfer of electrons from the anode to the cathode results in the generation of an electrical current.
[AU:Edits OK?]
©Bob Crimi
Development of pest resistance to insecticidal
transgenic crops remains a major concern
worldwide
1
.Although the use of Bt (Bacillus
thuringiensis) cotton and corn in the United
States alone constitutes 20–30% of all cotton
and corn grown there (totaling over 20 mil-
lion acres), almost all of these crops contain
only one of two Bt proteins for Lepidoptera
control (Cry1Ab for Bt corn and Cry1Ac for
Bt cotton). As a result, development of insect
resistance to toxins is a major concern, and
scientists continue to search for additional
genes to express along with either Bt gene
described above, or alone in plant cultivars
that can be planted either in a mosaic or
sequential fashion with Bt crops. [A
U: mean-
ing correct?] In this issue, Liu and coworkers
report the successful expression of a non-Bt
insecticidal protein in plants at high levels,
conferring activity against two different
orders of insects (Lepidoptera and
Coleoptera)
2
.These results have the potential
to help decrease the rate of insect resistance
development to toxins expressed by commer-
cially available insect-resistant transgenic
plants, such as Bt cotton and corn. In addi-
tion, expressing one protein with activity
against insects from two different orders
should allow increased control of insect pests
and reduce the time needed for making a
transgenic plant with such enhanced activity.
[A
U: paragraph edit OK?]
Thus far, scientists have been successful in
finding several new Bt proteins for these pur-
poses. Cotton containing both Cry1Ac and
Breathing new life into insect-
resistant plants
William J Moar
Expressing insecticidal proteins from sources other than Bacillus
thuringiensis in crop plants should reduce the likelihood for development of
insect resistance to toxins.[AU: shortened description OK?]
[AU:
Please cite Figure 1 within text]
William J. Moar is in the Department of
Entomology and Plant Pathology, 301 Funchess
Hall, Auburn University, Auburn, Alabama
36849, USA.
e-mail: moarwil@auburn.edu