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particularly controversial, however, is its
unclear relation to the thermodynamic basis
of plant water relations. Thermodynamics
dictate that since water must always flow,
energetically speaking, 'down hill', then the
water in the xylem elements must have
some property that reduces its 0 to at least
slightly lower than the 0 of the water in the
soil. Numerous experiments have indicated
that xylem water contains very few dis-
solved solutes (dissolved solutes are one
way 0 can be reduced), so a hydrostatic ten-
sion appears the only other alternative.
An additional complication in this
debate is that the measurements of little or
no tension in xylems have also been used to
question the validity of measurements made
with a device that is widely used to estimate
tension in the xylem of plants, known as a
pressure chamber I3. This side of the debate
has given rise to some novel experimental ap-
proaches, one being the spinning of a length
of branch about its center to create a known
tension based on centrifugal forces14. The re-
sults of such experiments were a reasonable
correspondence between the pressure cham-
ber measurements and the tension expected
from centrifugal forces, to a substantial ten-
sion of about -2 MPa (equivalent to the ten-
sion in a 200 m tall column of water). Hence,
it would appear that tensions can be created
in xylem vessels and measured accurately
by the pressure chamber. Some of the debate
surrounding the pressure chamber device
may stem from the initial belief that it gave
a direct measure of the tension in the xylem
per se 13, whereas a more accurate descrip-
tion is that it measures the matric compo-
nent of 0 in the leaf apoplast15,16,which may
not be equivalent to the tension in the
xylem under all circumstances.
The ongoing debate
So how does water get to the top of tall trees,
or for that matter, even short ones in drying
soil? If it is by tension-cohesion, then how
is the system engineered to avoid all the
technical difficulties of reliably transporting
metastable water, sometimes under harsh
environmental conditions? The question of
whether the transport is by a strictly thermo-
dynamic mechanism of water flowing down
a gradient in potential energy, or by an in-
teresting hybrid mechanism involving some
as yet unrecognized biological processes, is at
the heart of the current debate. I suspect that
as long as the controversy does not extend
itself to plant consciousness, raising doubts
about the plant's ability to transport water,
our food and fibre supply should be safe!
Acknowledgements
K.S. is grateful for the sabbatical support of
the CRC/Plant Science, the helpful com-
ments of J. Passioura and R. Munns, and
the lively discussions of these issues by the
Pomology Dept's Friday Regular Open
Graduate Section (FROGS).
Ken Shackel
Dept of Pomology, University of CNifornia, Davis,
CA 95616, USA
References
I Klein, H.A. (1988) The Science of
Measurement: a Historical Survey, Dover
2 Hales, S. (1727) Vegetable Staticks, or, an
Account of some Statical Experiments on the
Sap in Vegetables, Oldbourne
3 Dixon, H.H. and Joly, J. (1895) On the ascent
of sap, Philos. Trans. R. Soc. London Ser. B
186, 563-576
4 Nobel, P.S. (1991) Physicochemical and
Environmental Plant Physiology, Academic
Press
5 Hayward, A.T.J. (1971) Negative pressure in
liquids: can it be harnessed to serve man?
Am. Sci. 59, 434-443
6 Zimmermann, U. et al. (1994) Xylem
water transport: is the available evidence
consistent with the cohesion theory? Plant
Cell Environ. 17, 1169-1181
7 Smith, A.M. (1991) Negative pressure
generated by octopus suckers: a study of the
tensile strength of water in nature, J. Exp.
Biol. 157, 257-271
8 Balling, A. and Zimmermann, U. (1990)
Comparative measurements of the xylem
pressure of Nicotiana plants by means of the
pressure bomb and pressure probe, Planta
182, 325-338
Passioura, J.B. (1991) An impasse in plant
water relations? Bot. Acta 104, 405-411
Hayward, A.T.J. (1970) New law for liquids:
don't snap, stretch! New Sci. 45, 196-199
Balling, A. et al. (1988) Direct measurement
of negative pressure in artificial-biological
systems, Naturwissenschaflen 75,409-411
Canny, M.J. A new theory for the ascent of
sap - cohesion supported by tissue pressure,
Ann. Bot. 75, 343-357
Scholander, P.F., Bradstreet, E.D. and
Hemmingsen, E.A. (1965) Sap pressures in
vascular plants, Science 148, 339-346
Holbrook, N.M., Burns, M.J. and Field, C.B.
(1995) Negative xylem pressures in plants: a
test of the balancing-pressure technique,
Science 270, 1193-1194
Tyree, M.T. and Hammel, H.T. (1972) The
measurement of the turgor pressure and the
water relations of plants by the pressure-
bomb technique, J. Exp. Bet. 23, 267-282
Passioura, J.B. (1980) The meaning ofmatric
potential, J. Exp. Bot. 31, 1161-1169
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16
The gene-for-gene relationship: from
enigma to exploitation
Genetics has transformed plant pathology
on three occasions: at the turn of the cen-
tnry, when disease resistance was observed
in plants as yet another Mendelian trait; in
the mid-1940s, when H.H. Flor proposed
the gene-for-gene hypothesis to explain the
correspondence of disease resistance with
Mendelian traits (so-called avirulence
genes) in the parasite; and in recent years,
when several research groups first used
molecular genetics to identify and decode
the DNA sequences of plant genes required
for disease resistance. It was auspicious
timing, therefore, that the 1995 president
of the British Society for Plant Pathology
(BSPP), Ian Crute (HRI-Wellesbourne),
selected a theme for the society's annual
meeting entitled, 'The gene-for-gene rela-
tionship: from enigma to exploitation'*.
The meeting proved to be of wide inter-
national appeal, with more than 200
participants from around the world - nearly
double the usual attendance at BSPP
meetings.
Twenty-six speakers were assembled to
present the results of their research on the
genetics of a wide range of host and para-
site interactions. Ian Crute set the stage
with a presidential address in which he
drew on examples from his own career
(transcript to be published in Plant
Pathology). He was responsible for pioneer-
ing work on the genetics of host and para-
site genes involved in lettuce downy
mildew, with the result that it has become
one of the best genetically defined exam-
ples of plant parasitic symbiosis. Practical
*BSPP Presidential Meeting, University of
Warwick, UK, 12-15 December 1995.
benefits resulting from his work were
demonstrated by a description of how let-
tuce breeders and commercial growers can
now effectively manage the 'boom and bust
cycles' caused by continual adaptation of
the parasite to resistant cultivars and the
fungicide metalaxyl. A further recent con-
tribution he has made has been to establish
downy mildew in Arabidopsis as a prime
model for molecular genetic analyses of
host-parasite recognition.
Many of the talks were devoted to genetic
analyses of host resistance. Some of the
topics included: fine structural analyses of
regions in plant genomes containing a
complex of parasite recognition genes; mu-
tatienal dissection of signal transduetion
for primary host defence; and molecular
characterisation of genes required for
disease resistance. The genetic analysis of
1 O6 April 1996, Vol. 1, No. 4 © 1996 Elsevier Science Ltd