Effects of Electrostatic Fields, Microwaves and Electric Currents on Plants

The Effect of Externally Applied Electrostatic Fields, Microwave Radiation and Electric
Currents on Plants and Other Organisms, with Special Reference to Weed Control
Author(s): M. F. Diprose, F. A. Benson and A. J. Willis
Reviewed work(s):
Source: Botanical Review, Vol. 50, No. 2 (Apr. - Jun., 1984), pp. 171-223
Published by: Springer on behalf of New York Botanical Garden Press
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THE
BOTANICAL
VOL.
50
REVIEW
APRIL-JUNE, 1984
No. 2
The Effect of Externally Applied Electrostatic Fields,

Microwave Radiation and Electric Currents on
Plants and other Organisms, with
Special Reference to Weed Control
M. F. DIPROSE, F. A. BENSON
Departmentof Electronicand ElectricalEngineering,Universityof Sheffield
MappinStreet,Sheffield,SI 3JD UnitedKingdom
AND
A. J. WILLIS
Departmentof Botany, Universityof Sheffield,WesternBank
Sheffield,S1O2TN UnitedKingdom
I.
II.
III.
IV.
Abstract .----------------------------------------172
Sommaire.------------------------------------------------173
Introduction ..14...
..............
174
....
Definition of Forms of Electrical Energy
..
.
175
A. Electrostatic Fields .--------------.----.--..--..-----175
.
B. Microwave Radiation
.176
C. Electrical Discharges and Direct Electric Shocks
.
177
Electrostatic Fields and their Lethal Effects on Plants ..-------------------------------------------177
A. Introduction ..----------------------------------------------------------------177
B. Plant Growth in the Presence of Electric Fields
.....
178
C. The Lethal Effects of Electric Fields on Plants ..
179
.
D . C onclusions
186
..18------------------------------------------------------------------------------------------------------(i) Summary
186
(ii) Practical Weed Control by High Electric Fields ........................................ ...... 187
Microwaves and Weed Control ...---------------------------------187
A. Uses of Microwaves in Agriculture ---------------------------------........-...............
187
B. Laboratory Experiments with Microwave Radiation on Plants and Seeds
190
Copies of this issue [50(2)] may be obtained from: Scientific Publications Office,
The New York Botanical Garden, Bronx, NY 10458 USA. PRICE (Includes postage
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The Botanical Review 50: 171-223, April-June, 1984

? 1984 The New York Botanical Garden
171

172
THE BOTANICAL REVIEW
................................................
196
C. Field Experiments with Microwaves for Weed Control
D. The Effect of Microwave Radiation upon Soil Microorganisms and
N em ato des ..---- --- -- --- --- -- --- --- -- --- --- ----- --- --- -- --- ---------- --- 202
E. Conclusions ..204------................. 204
.
(i) Summary.------------------------------------------204
----206
(ii) Thermal or Non-thermal Mechanisms of Death.
207
(iii) Practical Weed Control by Microwaves .................
209
V. The Effect of Electric Currents Applied Directly to Plants and Soils .
209
.
A. Soils, Plants and Applied Currents
.....
212
B. Weed Control by High Voltages ...........
215
C. Conclusions .............................................................
(i) Summary.----------------------------------------------------215
(ii) Practical Weed Control by Electric Discharges and Currents.
...... 216
217
VI. Acknowledgments
................
217
VII. Literature Cited.------------------..----------------------------------------------------------------------------
Abstract
A wide-ranging review is presented of the effects of various forms of
externally applied electrical energy upon plants and other organisms. Although investigations involving both small and large amounts of energy
directed at the targets are considered, a particular emphasis of this review
is the feasibility of each type of electrical stimulation for weed control.
Electrostatic fields ranging from 100 V m-I to 800 kV m-I have been
applied to plants under laboratory conditions and in field trials since the
1880's. Some beneficial effects have been reported (e.g. increase in yield
from both cereal and vegetable crops), but the results have been erratic
and the electrical conditions leading to definite benefits on a large scale
could not be confidently predicted from early studies. High electric fields
are reported to damage plants if currents greater than 10-6 A are induced
to flow through leaves causing corona discharges from the tips. The nature
of the damage and the effects on metabolic processes are discussed. The
results from experiments on the growth of plants in which the density
and charge of air ions have been varied are also reviewed.
The effects of microwave radiation (mostly 2450 MHz) upon seeds,
plants and other organisms in soil are discussed. These effects depend
upon the power density of the radiation and the electrical properties of
the targets. Factors such as size of seeds and plants, shape and moisture
content are important, as are the properties of the soil irradiated (notably
water content). Although microwaves can be effective in killing plants
and also seeds that are buried several centimeters deep in soil, high power
equipment is required and treatment times are long e.g. a 60 kW machine
could take up to 92.6 hours per hectare. Other experiments reported show
that microwave radiation can kill nematodes in the soil and that it is also
very effective in killing fungi and bacteria. The potential of the various
possible uses of microwave radiation in agriculture is also described.

ELECTROSTATIC FIELDS, MICROWAVE RADIATION
173
Electric currentshave been caused to flow through plants by the application of electrodesto the leaves. The effectsrangefrom nil, when 50100 V and 1 or 2 ,uAare used, to very strikingwhen voltages from 5 to
15 kV areappliedcausingcurrentsof severalamperesto flow and resulting
in the rapiddestructionof the target.Smallelectriccurrentspassedthrough
soil containingplants are reportedto increasetheir growth.The effectsof
small currenton the growth of individual leaves are reviewed. The use
of high voltage tractor-borneequipment for weed control is also considered.
Sommaire
Une revue a largesthemes presenteles effets des diverses formes d'application externed'energieelectriquesur les plantes et autresorganismes.
Bien que des recherchescomportanta la fois de petiteset grandesquantites
d'energiedirigeessur les cibles en question soient prises en consideration,
un des aspects particuliersde cette revue est la possibilite d'application
de chaquetype de stimulationelectriqueau controledes mauvaisesherbes.
Depuis environ 1880, les plantes ont ete soumises, soit en laboratoire,
soit lors d'essais sur le terrain, a des champs electrostatiquesallant de
100 V m' a 800 kV m'. Quelqueseffets benefiquesont ete enregistres,
par exemple, l'accroissementde la productiondes recoltes de cerealeset
de legumes; mais les resultats etaient irregulierset les conditions electriquesconduisanta des profitsbien determinesa grandeechelle ne pouvaient pas etre predites avec confiance des premieres etudes. On s'est
apercuque de grandschamps electriquespouvaient deteriorerles plantes
si des courants superieursa 10-6 A etaient amenes a circuler dans les
feuilles, entrainantdes dechargesde la couronne a partirdes pointes. La
nature des degats ainsi que les effets sur les procedes metaboliques sont
ici etudies, de meme que les resultats des experiences sur la croissance
des plantes pour lesquelles la densite et la chargedes ions dans I'air ont
ete changes.
Les effets du rayonnementpar micro-ondes (pour la plupartde 2450
MHz) sur les graines, les plantes et autres organismes du sol sont ici
exposes. Ces effets dependentde la densite electriquedu rayonnementet
des proprieteselectriquesdes objectifs. Des facteurstels que la taille des
graineset des plantes, leur forme et leur teneur en humidite sont importants, comme le sont les proprietesd'un sol irradie(notammentsa teneur
en eau). Bien que les micro-ondes puissent etre efficaces pour tuer des
plantes et aussi des graines enterreesa plusieurs centimetres de profondeur, un important appareillageelectriqueest necessaireet les temps de
traitementsont longs; par exemple une machine de 60 kW peut prendre
jusqu'a 92,6 h ha-'. D'autres experiencesdemontrerentque le rayonnement par micro-ondes pouvait tuer les nematodes dans la terre et qu'il

174
etait tres efficacepour detruireles champignonset les bacteriesdu soussol. Le potentiel des diverses utilisations possibles de la radiation par
micro-ondes en agricultureest egalementdecrit.
Des courantselectriquesont ete amenes a circulera traversdes plantes
par l'applicationd'electrodessur les feuilles. Les effets, nuls quand 50 a
100 V et 1 a 2 ,uA sont utilises, sont par contre frappantsquand des
voltagesde 5 a 15 kV sont appliques,entrainantla circulationde courants
de plusieurs amp'ereset la destruction rapide des cibles. On remarque
cependantque, de petits courantselectriquesenvoyes dans un sol contenant des plantes, accelerentleur croissance;les effets d'un faible courant
surla croissancedes feuillesindividuellessont ici reexamines.L'utilisation
d'un appareillagea haut voltage mobile pour le controle des mauvaises
herbes est aussi pris en consideration.
I. Introduction
Mechanical methods of removing weeds, e.g. hoeing and tilling, have

been practicedfor centuries(Crofts, 1975). The use of chemical methods
for weed control has been expandingat a fast rate since the mid 1940's
(House, 1967). In recent years electricalmethods have been investigated
in this connection, some of which promise to be of practicalvalue. This
review discusses the control of weeds, and also of plant growth more
generally,by means of such methods.
Weeds can be subjectedto electricalenergyby use of electrostaticfields,
microwaves, electric dischargesor direct electric shocks using either alternating current (a.c.) or direct current (d.c.). These techniques have
several advantagesover present methods of weed control and, although
they cannot be regardedas a panacea for all problems, electrical means
appear to offer viable and useful additions to the stock of weed control
methods.
Duringand followingthe applicationof an electricshock or microwave
or laser irradiation,the energy,rapidlyabsorbedby the "load," is largely
dissipated as heat in the plant. No residue is left to contaminatethe soil,
an importantfeaturein view of rising concern about the contributionby
herbicidesto environmentalpollution. The electricalenergy can also be
directed to where it is required,even under fairly adverse weather conditions. Electrodescarryingthe currents,or the radiation being directed
towards the ground are, unlike sprayedherbicides, not blown by winds,
so areas can be treated under a wider range of weather conditions than
previously possible. Microwave radiation penetratesthe soil to a depth
of several centimetersdependingupon the appliedpower,its wavelength,
and the composition and moisture content of the soil. This microwave
treatment may prove better than the use of flame guns for destroying

ELECTROSTATIC
FIELDS,MICROWAVERADIATION
175
weed seeds of the type that can withstandhigh temperatures,e.g. as great

as 1270Cif slightly below the surfaceor if in cracksin the soil (Sampson
and Parker, 1930). Microwave power would reach these seeds and still
be as effective in killing them as if they were on the surface.
While electricalmethods of weed control are a relatively recent innovation, some forms of electrical and electromagneticenergy have long
been known to be essential in agriculture-notably electromagneticradiation in the visible wavelengthregion (Borthwick, 1965) necessaryfor
photosynthesis. Other plant processes in which light is involved include
photoperiodismand germination.Furthermore,higherfrequencyelectromagnetic waves such as y-rays may be important in evolution, leading
to the formationof new species, by causingmutationswhen absorbede.g.
by seeds (Nelson, 1965).
The naturalelectricalstate of the atmospheremay be importantto plant
growth (Wheaton, 1970). The air contains both positive and negative
ions, continuouslyprovidedby radioactivedecayof elementsin the earth's
surfaceand some cosmic radiation (Chalmers, 1967). Ion densities vary
throughoutthe day but over open land are in the order of 1.5-4.0 x 103
ions cm-3. Kotakaand Krueger(1968) have examinedthe effectsof varying ion densities of air on the growth of barley, oats and lettuce. They
reportthat increasesin ion density lead to vigorous, acceleratedgrowth.
Alternativelyplants exposed to ion-depletedatmosphereslack vigor and
have soft leaves (Kruegeret al., 1965).
The effects of electricalenergy are thereforeimportant to plant life in
many ways and electromagneticradiationis vital to it. For weed control,
however, largerthan normal applications of electrostaticfields, electric
discharges,and microwave radiationto plants are necessary.These treatments are reviewed below, following a short explanationof the technical
definitions of the various forms of electricalenergy.
II. Definition of Forms of Electrical Energy
Small electric potentials exist across the membranesof plant cells and
within whole plants. These can give rise to potentials of 200-300 mV
betweendifferentpartsof the plant (Lund, 1947). Suchnaturallyoccurring
phenomenaare not consideredin this review which deals with the various
ways in which plantscan be subjectedto electricalenergyfrom an external
source-typically many orders of magnitudegreaterthan internallygenerated electricaleffects.
II.A ELECTROSTATICFIELDS
An electrostatic field is an electric field that is static in time, i.e. its

strengthdoes not vary throughtime (Nussbaum, 1966). The field can be

176
THE BOTANICALREVIEW
formed by placing two conductors apart from each other (they need not
necessarilybe parallelor both planar)and connecting them to a voltage
source. The electric field strength for parallel planar conductors is defined as
V
E= d
where E is the electric field strengthin the air gap between the electrodes
(conductors)measured in volts per meter, V the voltage applied to the
conductorsin volts, and d the distance between them in meters. For the
parallel-plateconfigurationthe field is uniform between the conductors,
but differentexpressions exist to describe other more complex non-uniform field patternsthat arise, for instance, with one planarelectrode(e.g.
earth) and an overhead wire, or when a plant is introducedbetween two
parallelplates.
II.B MICROWAVE RADIATION
Microwaveradiationis the name given to electromagneticradiationof

which the wavelengthsare much greaterthan those of light, i.e. they are
measuredin mm or cm (Glazierand Lamont, 1958). A wavelengthcommonly used in weed controlis 12.25 cm which correspondsto a frequency
of 2450 MHz. Most of the papersreferredto in this review that describe
microwaveexperimentsuse this frequency.This is one of the frequencies
allocated, by internationalagreement, to microwave power devices for
domestic and industrialapplications.A wide rangeof equipmentis commercially available which produces the necessaryradiation powers. The
power absorbed, Pabs, by an irradiated dielectric load which is converted
to heat is given by
Pabs = 0.556fE2Er"x 10-10 Wm-3
where f is the frequencyin Hz; E the field strengthof radiation at point

of absorptionin Vm- 1;Er" is the relative dielectricloss factor(Nelson and
Wolff, 1964; White, 1970).
Thus the power absorbeddepends upon several factorsincludingproperties of the load itself such as the relative dielectric loss factor, and the
homogeneity of the medium. The electric field strengthat any point in
the load depends upon the load shape and the dielectricpropertiesof the
medium, as well as the strengthand frequencyof the irradiatingfields.
The radiationpenetratesall parts of the load simultaneouslywhich is in
contrastto the normal heating process where heat is conductedfrom the
outside of the load towards the center. The speed of action is one of the
main benefits of microwave radiation.

ELECTROSTATIC
177
II.C ELECTRICAL DISCHARGES AND DIRECT ELECTRIC SHOCKS
Electricdischargesin the context of this review are taken to mean the

dischargeof a quantity of electricity into the plant from a high voltage
electricalenergydevice, usually, but not necessarily,a capacitor(Duffin,
1965). When the electric chargeis pulsed, the generatorhas to chargeup
a capacitor,and when that operationis completedthe energyis transferred
to the load. Repetition rates depend upon the generatorsize, the storage
capacitanceand the propertiesof the load under discharge.Russian researchershave used this method with up to 60 kV pulses of 1 Asduration.
The dischargeelectrodesneed not necessarilytouch the load if very high
voltages are used, since the dischargecan jump across an air gap. Alternatively, a pulsed high voltage discharge can be applied from the secondarywindingsof a high voltage transformer,without using a capacitor,
by a suitable control of the input voltage to the primarywindings.
Direct electric treatmentis a continuous process whereby a generator
is physicallyconnectedto the plant by two electrodes,or by one electrode,
the groundbeing used as the other. The voltage can rangefrom 500 V to
several kilovolts as required,and can be a.c. or d.c. The current,I (A),
flowingthroughthe plant is determinedby its electricalresistance,R (Q),
and the applied voltage, V (volts), so that
V
R
The units of alternatingvoltage and currentare volts or amps r.m.s. The
latterterm standsfor "root mean square"and describesthe effectivevalue
of a voltage or currentwaveform that varies continually with time (e.g.
an alternatingcurrentwhich has the same effect on a load as a 1 A direct
currenthas a value of 1 A rms).
III. Electrostatic Fields and their Lethal Effects on Plants
III.A INTRODUCTION
A number of studies of the effects of electric fields on the growth of a

range of plants have been made. The Great Plains area of the United
States sometimes suffersfrom severe dust storms, in which the normal
electrical balance of the atmosphere is considerablydisturbed and very
high electric field strengths exist near the ground level. Miller (1938)
reports that, after these storms, damage observed in some of the crops
was attributedto electrical action. Shlanta and Moore (1972) have also
observeddamage(brownedleaf tips) in the "natural"grassof a mountain
meadow in New Mexico after storms. Such observationshave led to the
investigationof the possibilities of a "lethalelectrotropism"in plantsand

178
THE BOTANICALREVIEW
to the demonstrationsthat growth can be stopped and leaves and whole

plants killed by higher electric fields.
III.B
PLANT GROWTH IN THE PRESENCE OF ELECTRIC FIELDS
Earlyresearchsuggestedthat the effects of subjectingplants to electric

fields were beneficial. This topic has been the subject of research and
debatefor many years(Ellisand Turner,1978;Pohl, 1977;Sidaway,1975;
Wheaton, 1970), but the beneficial effect is not now generallyaccepted.
From the mid eighteenthto the earlytwentiethcentury,findingsindicated
that increasedyields from both cereal and vegetable crops could be obtained by applying electrostaticfields to plants while they were growing
(Hendrick, 1918; J0rgensen and Priestley, 1914; J0rgensen and Stiles,
1917; Newman, 1911; Shibusawaand Shibata, 1930).
Between 1885 and 1903 Lemstrom (1904) undertook various experiments on the effects of electrical fields on plants in different places in
Europe-from Finlandand Swedenin the north,to Englandand Burgundy
in the south. His results,while encouraging,were not very consistent (one
factorbeing the unreliabilityof earlyhigh voltage equipment).His general
conclusion was that electricfields, applied to crops by wires strungabove
the growingareas,werebeneficial,producinghealthyplantswith increased
yields. This applied to root crops, vegetables, cereals and strawberries.
He proposed that the best times for applyingthe electrostaticfield were
for four hours in the early morning and another four during the late
afternoon.The electricitycould be applied all day duringcloudy weather
and during nights of moist weather. Application during dry conditions
and in strong sunshine, however, could have adverse effects.
Lemstrom'sstudies were extended by several workers.In general,the
same encouragingresults were obtained. Blackman's (1924) field trials
with cereals and clover-hay between 1917 and 1920 produced 18 sets of
results of which 14 showed increasedyields; 9 of these increaseswere of
at least 30%.In pot-cultureexperimentscarriedout on maize, wheat and
barley, the average yield increase, when these cereals were subjectedto
weak electric currents,was over 11%(Blackmanand Legg, 1924). The
Board of Agricultureand Fisheries of the U.K. even establisheda Committee for Electroculturein 1918;however, its finalreportin 1937 (Board
of Agricultureand Fisheries, 1918-1937) stated that the results of 18
years of researchwere not conclusive and it was not possible to predict
confidentlythe benefits of electricaltreatmentsto crops.
The encouragingresults of the Europeanworkerswere not, however,
shared by their American colleagues, who, in two fairly comprehensive
sets of controlled trials, found no significantbeneficialeffects of electric
fields on growth(Briggset al., 1926; Collins et al., 1929). In view of these

ELECTROSTATIC
179
findingsand the inconsistency of other results, the use of electricaltreatments of plants lost favor. Apart from a few supporters(Sidaway, 1969;
Sidawayand Asprey, 1968), the methods did not attractattentionalthough
more recently interest has been shown in the effects of air ions, rather
than electric fields, on plant growth (Bachmanet al., 1971; Kotaka and
Krueger,1968; Kotaka et al., 1965; Kruegeret al., 1965, 1978).
III.C THE LETHAL EFFECTS OF ELECTRIC FIELDS ON PLANTS
The term "lethal electrotropism"was suggestedby Murr (1964a) following a series of experiments on the effects of electric fields on plants.
In these experiments(Murr,1963a, 1963b) an electricfieldwas established
between two aluminium wire grids (0.24 m2). A lower electrode was
situatedbelow the soil in a plot in which seedlingsof orchardgrass(Dactylis glomerata)were planted. An upper electrode was suspended above
the soil and adjustedin height to vary the electric field strength,but was
never more than 10 cm above the tops of the plants. Temperatureand
light intensity were controlled(unspecified)and a 16-hourday lengthwas
used. The controlplots had the same electrodearrangementas the "active"
ones, but without the voltages applied. The top electrodewas made positive and the bottom one connected to the negative of the power supply
simulatingthe earth's naturalelectric field (Chalmers, 1967).
Murr(1963a, 1963b) observed that duringcontinuous exposureto the
electrostaticfieldsthe leaf tips of the seedlingsbeganto brown,as if burnt,
and he noted the similarityto mineraldeficiencysymptoms. Betweenthe
region of leaf tip "burning"and the normal tissue was a small strip of a
much deeper green color than usual. He also found that damage spread
downwardsfrom the tip at a fasterratethan the growthof the plant (Murr,
1963b). The plants were clipped to a height of 2.5 cm after two weeks,
twice more at weekly intervals, and the dry weights of the clippings obtained. Murrdefineda damage factoras the proportionof the dry weight
of electrifiedto control samples (averageof three results) expressedas a
percentage.For orchardgrass, damagewas found to rise to 25%at a field
strengthof 50 kV m-1 and then rapidlyincreasedto 50%at 75 kV m-l.
Similarresultswere obtainedwith seedlingsof reed canarygrass(Phalaris
arundinacea)(Murr, 1963b). Transversesections of some leaves showed
that the epidermalcells had been destroyedin the brownedtip and damaged in the darkgreenzones. Therewas complete absenceof cell structure
in the tip area and chloroplastderangementin the dark green band.
Murr believed a possible cause of this damage was the migration of
ionized salts to the leaf tip under the action of the electric field. The
resultingconcentrationunbalancemight upset normal osmotic phenomena and cause ruptureof the cells. To investigate this furtherhe applied

THE BOTANICALREVIEW
180
Table I
The concentrationof elementsin the leaf tips of orchardgrass(Dactylisglomerata)
seedlingsafter exposureto electrostaticfields (afterMurr, 1964b)
Electrostatic
strfielgth
(kV m-')
Phosphorus
Nitrogen
30
50
104
94
95
97
75
97
98
Iron
Zinc
Aluminum
(percentage of control samples)

119
177
104
169
202
258
317
233
324
field strengths of 30, 50 and 75 kV m-1 to seedlings of orchard grass (in

the same experimental regime as before) and took clippings at 2, 3, 4 and
5 weeks (Murr, 1964b). Dried samples were used for mass spectrometric
and micro-Kjeldahl analysis. The results were contrary to expectation,
and there were no significant differences in the quantities of phosphorus,
nitrogen, calcium, magnesium or potassium between the electrified and
control samples. The minor elements iron, zinc and aluminum did show
increases, however, which ranged from 104% to 324% (Table I).
Murr (1 964a, 1964b) concluded that the lethal damage was not caused
by the drift of the ionized salts, leading to the bursting of cells and dehydration, but suggested that metabolism was accelerated and a "general
tissue deterioration" ensued. He regarded (1964b) the increase of iron,
zinc and aluminum in the leaf tips of exposed plants to be associated with
accelerated "metallo-enzyme activity" affecting respiration, ultimately
leading to tissue destruction. He also interpreted changes in the density
of chloroplasts as evidence for "metabolic acceleration" (Murr, 1964a).
However, while the highest densities of chloroplasts, in the tips of leaves
of orchard grass damaged by exposure to an electrostatic field of 40 kV
m- I (Murr, 1964a), were found in the deep green regions (below the brown
tip), even here the chloroplast density was lower than that in untreated
leaves. The deep green of treated leaves was suggested (Murr, 1964b) as
attributable to the effect of traces of ozone in oxidizing porphyrin groups,
electrostatic fields leading to an increase of porphyrin.
The deep green color of leaves of plants exposed to static electric fields
had been reported by several earlier investigators. Priestley (1907) noted
that young blades of wheat from electrified plots were, in the opinion of
many observers, darker green than the control plants. In continuation of
his experiments, Priestley (1910) again remarked on the color difference,
reporting that other workers had observed a darker green, this being
especially noticeable in wheat. Blackman and J0rgensen (1917), experimenting with oats, reported that, after one month, the electrified crop

181
was taller and greenerthan oats not subjectedto electric fields. Again in
an experiment at Rothamsted ExperimentalStation in 1917, Blackman
(1924) found that barleywas tallerand greenerafter20 days of application
of the electric field than that in the neighboringcontrol plot, althoughthe
visual differencebetween the crops subsequentlybecame less marked.
Priestley( 19 10) suggested(withoutproof)that the deepergreenin wheat
might result from a slight but continuous amount of nitratesbeing added
to the soil by the overheaddischarge,perhapsformedin a similarmanner
as the combination of oxygen and nitrogen producedby thunderstorms
is washed into the soil by rain. One soil test (Priestley, 1907) indicated
some three times the amount of nitrogenin the soil beneath an overhead
discharge than in the soil in the control plots, but unfortunatelymeasurementswere taken only after the crops had been harvested.However
Blackman(1924) reportedno appreciabledifferencesin soil nitrogencontent before and after the application of electric fields to oats. A further
unconfirmedsuggestionby Priestley(1907) is that plants exposed to electrostaticfields may utilize atmosphericnitrogendirectly,perhapsby combination of gaseous nitrogen with carbohydrateswithin the plant.
Hart and Schottenfeld(1979) also observed that leaves of pole beans
(Phaseolusmultiflorus)becamedarkergreenwhen exposed to electrostatic
fields resulting in corona current from a few points on the leaf edges.
Prolongedexposurecaused loss of turgorand collapse of the plants. The
pole bean plants were grownin individual containers(20?C;30%relative
humidity). Soil moisture content was monitored by measuringthe resistance between two brass posts set 4.5 cm apart.The plants were 5-10 cm
high at the time of treatment(estimatedfrom Fig. 1. of Hart and Schottenfield, 1979), and the mesh electrode was about 5 cm above the top of
the plant. With the electrode chargednegatively, plants sustained 2 ,uA
of corona currentfor 8 hours with no visible effects, and with 20 AA of
corona currentthere was little change in soil resistanceafter 5 hours but
the plant began to droop and one leaf became dark green and lost "texture."At 50 ,A, however, leaf discolorationdeveloped rapidlyand stem
collapseoccurredwithin 45 minutes.Withthe electrodechargedpositively
and 20 ,A of corona current,wilting occurredmore rapidlyand therewas
a largeincreasein soil resistance(about 35%in 3 hours).If slightlydrooping plants were waterednear their stems they became turgidagain. Hart
and Schottenfeldattributedthe effectsto severe waterloss from the leaves
caused by the corona current.
Bachmanand Reichmanis (1973a) applied various strengthsof electric
field to single leaves of barleythat were five days old and 5 cm long with
the cut end of the leaf in contactwith the negativeelectrode,and a positive
electrodesuspendedabove the leaf. Their resultsindicatedthat with fields
less than 40 kV m- I leaf tip burningwould not occur,but at a field strength

182
THE BOTANICALREVIEW
r damagewas almost instantaneous(afteronly a few seconds

of 400 kV mof exposure).Above 800 kV m- ' the density of positive air ions and level
of ozone were shown to build up very rapidly.The time of the appearance
of damagewas found by Bachmanand Reichmanis to be proportionalto
1/V2. They also investigatedthe speed of propagationof the damagedown
the leaf and found this to depend on the field strength. Although the
damage at the tip appearedvery quickly, the subsequentspreadwas at a
reducedrate, e.g. for 250 kV m-I damage appearedafter 10 seconds and
after 2 minutes it was about 0.8 mm down the leaf; after 60 minutes it
was only about 2 mm in extent. Similarly for 166 kV m-l the damage
appearedafter 20 seconds and after 2 minutes it was about 0.2 mm in
extent; after 60 minutes it was only about 1.1 mm down the leaf.
Damagein airwas comparedto that in nitrogenand hydrogenby passing
these gases over the leaf duringelectrification,and the steady progression
of destructionwas observed. The spread of damage was much higher in
the presence of hydrogen than in nitrogen or air, e.g. for 2 minutes'
exposurewith 10 kV voltage, 1 mm of damagedtip was observed in air,
and more than 4 mm in hydrogen.
From these results, Bachman and Reichmanis (1973a) concluded that
the damagewas probablycausedby glow and brushdischargesat the leafair interface, so they made subsequentobservations using a microscope
in a dark room. They observed an ivory-coloredglow around the tip of
the leaf and usually two purple-coloredbrush discharges,one on either
side of the leaf at the junction of the damaged and undamagedtissue.
These observations,in their view, confirmedthat damagewas caused by
glow discharges.This damage mechanism is substantiallydifferentfrom
that earlierproposed by Murr(1964b) of acceleratedrespirationcausing
membraneruptureand cell dehydration.The results also differ in other
respects.Murr(1963a, 1963b) reporteddamage at 40 kV m-l and below
and an intermediatezone of darkgreen,while Bachmanand Reichmanis
stated that no damage occurredbelow 40 kV m'-I and made no mention
of color changes.
After observing the physical characteristicsof the damage, Bachman
and Reichmanis (1973b) concentratedon the growthrates of plants subjected to electrostaticfields. By using both sloping and horizontal upper
electrodes,they found that barley seedlingsgrew until they were about 2
cm from the electrode and then stopped so that the plant tops took the
profile of the electrode. At this point the field strengthcorrespondedto
about 800 kV m- 1. Similarly,growthratesabove 200 kV m- I were found
to be inhibited when comparedwith controls, with 300 and 400 kV m- I
having strongeffects. By extrapolatingthese growthrate resultsthey predicted that a zero value should occurat approximately800 kV m- I which
agreedwith their earlierfindings.

183
Bankoskeet al. (1976) investigatedthe effectsof 60 Hz electromagnetic

fieldsupon plantsunderpowertransmissionlines. They carefullydesigned
their laboratoryexposurechamberto have an even fielddistributionwhen
empty and then calculatedand measuredthe effectof placingplantsinside.
The field pattern was disturbed considerably and enhanced in the immediate vicinity of the plants. Only with a dense growth of plants with
a uniform top surface could the electric field be defined as the applied
voltage divided by the distance between the top of the plants and the
electrode above. Single plants or small groups caused considerabledistortion but if the plant height was less than 25%of the electrode spacing
no more field enhancement occurred than for a single plant beneath a
transmission line. Plants with sharp-pointedleaves have a greaterconcentrationof electricfield aroundtheir tips and consequentlybreakdown
owing to corona currentat lower voltages than broad-leavedplants.
To define experimentalconditions Bankoskeet al. give "undisturbed"
field strengths,i.e. the field strengthwith an empty chamber,which bears
no direct relation to the actual field strengthsat the surfacesof leaves in
the chamber. This is also recognized by Hart and Schottenfeld (1979)
who preferredto state electrical exposure conditions in terms of AA of
corona currentratherthan unrepresentativefield strengthvalues.
Murr(1965a) uses a very simple model of a column of dielectricslabs
to representthe air, leaves, roots and soil between the electrodes.This is
unrealisticexcept in the one previously mentioned case of a very dense
growth of plants whose top surface forms a new ground plane. Most of
Murr'sexperimentsare with a single plant or groups of separatedplants
(e.g. Fig. 5, Murr, 1965a) and so his values for field strengthsare not
valid, especially his dynamic field strengthvalues.
Enhancementof fieldstrengthowingto the radiusof curvatureof objects
placed in electric fields means that it is impossible to relatethe responses
of differentspecies of plants, or even differentspecimens of one species,
to particularvalues of electrostaticfield strength.Each plant and part of
a plant will experiencedifferentelectricalforces at the leaf surfacewhen
placedin a uniformelectricfield. Comparisonbetweenexperimentsbased
on field strengthvalues is extremely difficult.
Bankoskeet al. (1976) determinedthat corona currentwas responsible
for leaf tip burningof corn (Zea mays) and alfalfa(Medicagosativa). The
alternatingfield (60 Hz) had an undisturbedvalue of 50 kV m-1 and
plants introduced into this had their uppermostleaf tips damaged after
exposurefor seven days. Coronadischargeswerephotographedat the tips
of leaves when plants were placed in "undisturbed"field strengthsof 2550 kV m-', with 22.5 kV m-1 as a damage inception level. Kentucky
bluegrass (Poa pratensis) showed leaf tip burning after a few days in
"undisturbed"field strengthsof 50 kV m-l, 60 Hz. The damagedid not

184
progress further down the leaves after one week and was 5-7 mm in
extent. At 25 kV m-l (60 Hz) damage was limited to 1-2 mm. When
placed in "undisturbed"electric field strengthsgreaterthan 30 kV m-'
leaves were seen to flutterowing to corona-inducedmotion.
Murr(1965b, 1966a, 1966b, 1966c)investigatedthe effectof "reversed"
(i.e. upperelectrodenegative)electrostaticfields,alternating(60 Hz) fields
and magnetic fields upon the growth of young plants. He concluded that
it was possible to stimulate their growth if the electricalconditions were
carefully chosen. If the electric fields became too large, corona current
flowedand causeddamageto leaf surfaces.The thresholdvalues per plant
for damagewere 5 x 10-7 A for orchardgrass(Dactylisglomerata)(Murr,
1965a), 1.5 x 10-8 A for sweet corn (Zea mays) and 3 x 10-8 A for wax
beans (Phaseolus vulgaris)(Murr, 1966c).
Murr proposed (1965b) a modification to his earlier theories (1963a,
1963b, 1964a, 1964b) to include the effect of corona current. It was
suggested that this caused damage to the epidermal layer of a leaf by
stimulating respiration and metabolism. Moderate electric fields led to
epidermaldamage and were believed to result in respiratoryaction sufficient to stimulategrowth but not to cause substantialleaf tissue damage.
Raising electric field strengths was thought to cause over-stimulation,
enzyme toxicity and sufficientepidermaldamageto resultin death of leaf
tissue. Murr(1966a) concluded that currentbelow 10-16 A per plant had
no effect but growth stimulation occurredin the range 10-I5 to 10-9 A
per plant. At 10-8 to 10-6 A per plant, leaf damage occurs, and plant or
leaf destructionoccurs at 10-5 A and above.
Blackman and Legg (1924), while examining the stimulation of the
growth of barley by electric fields, used currentsfrom 0.3 x 10-9 A to
175 x 10-9 A per plant to maximize the response. The higher currents,
however, were found to be injurious:above a level of 10-8 A per plant
there was damage to the plant tissue. Collins et al. (1929) claimed that
passing 75 x 10-9 A per plant had no effect on maize, but with larger
currentssome plants showed injury.
Scott (1967) reportedthat attempts to modify growthby using electrostatic fields were inconclusive, most having no effector retardinggrowth.
He also noted that lethal damagemay be causedby coronadischargefrom
leaf tips.
Kruegerand others (Kotaka and Krueger, 1968; Kruegeret al., 1978)
have experimentedwith changes in the density of air ions. It is believed
that the effects observed in the past may have been due more to the
atmospheric ion conditions that the electric fields per se. They showed
that increasingthe air ion density (from the normal values of approximately 4.0 x 103 ions cm-3 to approximately3.5 x 107 ions cm-3) resultedin speededup respirationand growthratesof youngbarleyseedlings

ELECTROSTATIC
185
(Kotakaet al., 1965) as comparedto controls. Increasesin air ion density

have also been reportedto increase the chlorophyllcontent slightly and
the cytochrome content considerably in treated plants (Kruegeret al.,
1963), to speed up chlorosis of plants grown in iron-depleted environments (Kotakaand Krueger,1968; Kruegeret al., 1964), and to stimulate
ATP metabolism (Kotaka et al., 1968) in isolated spinach chloroplasts.
Complementingthese results, Kruegeret al. (1965) found that barley
seedlingsgrownin ion-freeatmospheresshow retardedgrowth,lack rigidity and have soft leaves. Blackmanand Legg(1924) and Hicks (1957) have
experimented with plants and trees in areas surroundedby wire mesh.
All reportedretardedgrowth,lack of turgorand soft leaves. A wire mesh
surroundingplantswould act like a Faradaycage so the atmosphereinside
it would be relatively free of electric fields and air ions. Kruegeret al.
(1978) exposed barley seedlings (Hordeumvulgarevar. CaliforniaMariout) to electric fields in an air ion-free chamber and to electric fields in
an air ion-enriched environment (1.7 x 105 small negative ions cm-3;
currentaveraged 10-11 A per plant). They found an increase in growth
rate of the ion-treated plants compared with those in an electric field
alone, but no differencebetween the latterand control groupsgrownwith
an ion-free and no electric field regime. They concluded that the air ions
were biologicallyactive and responsiblefor the increasesin plant growth,
althoughthey were unable to distinguishwhetherthe effects were due to
the air ions alone or to their presence allowing electric currentsto flow
through the plant. They pointed out that many previous workers had
examined the responsesof plants when subjectedto electricfieldsbut had
not taken into account the presence and role of air ions.
Bachmanet al. (1971) found that under their experimentalconditions
ozone appearedwith currentsof 5 MAthrougha single leaf or 0.2 PA per
leaf if there were 20 plants. They recognizedthat the electric field at the
tip of a barley leaf would be much higher than that calculatedfrom the
separationof leaf and electrodeowing to the small radiusof curvatureof
the leaf tip. Also the edge of a barley leaf is covered with tiny "spikes"
whose diameteris about 1000 times smallerthan the leaf tip radius.Thus
very high electric fields would exist around the "spike" tips and corona
dischargeswill appearat much lower potentialsthan would be needed for
dischargesfrom the leaf tip (equivalentto a few hundredV m'-I between
planarelectrodes),so ozone will be producedand lethaldischargecurrents
can flow at quite moderate exposure levels.
Ozone can damage plants and is produced at ground level by point
dischargesduring thunderstorms.However, Shlanta and Moore (1972)
commentingon the leaf tip burningof grassconsiderthat it is the discharge
thatburnsthe grassratherthan the ozone. AlthoughBachmanet al. believe
that it may inhibit growthand Menser et al. (1963) reportozone damage

186
THE BOTANICALREVIEW
to tobacco leaves, complete death of a plant from the gas has not been
suggested.Bankoskeet al. (1976) measuredthe amounts of ozone present
when leaf tip burningoccurredduringcorona dischargefrom leaves. The
level was 16 parts per billion (ppb) compared to an ambient level of 8
ppb. It was concludedthat ozone was unlikelyto have causedany damage.
Blackmanet al. (1923) passed small currents(0.5 x 10-10 A per plant)
throughbarleyseedlings(Hordeumvulgarevar. Goldthorpe).The current
was generatedby means of a needle connected to a high voltage source
(ca. 10 kV) suspended 20 cm above the plants, applied with the needle
electrode positive for one hour or three hours and for three hours with
the needle negative. The positive polarity produceddefinite increasesin
growth rate over controls (no electric field or current)which increased
with time and persistedfor up to four hours afterthe currentwas stopped.
The negative polarityresultedin an initial increasefollowed by a steadily
decreasingrate of growthcomparedwith controls. When the currentwas
switched off, however, growth rates began to rise again still above the
control values but less than those of the positively chargedset. Further
workled them to concludethat neitherthe "electricwind" nor the gaseous
byproducts(i.e. ozone and nitrogenoxides) of the corona dischargewere
responsiblefor the effecton the growthof the plantswhen a currentpassed.
III.D CONCLUSIONS
III.D (i) Summary

Considerableeffort was expended in the first third of this century investigatingthe effectsof electric fields on the growthof plants (Jorgensen
and Priestley, 1914; Lemstrom, 1904; Newman, 1911). Although some
of the resultswerevery encouraging[e.g.field trialswith cerealsand clover
hay (Blackman, 1924) showed increases in yield of over 30% in half of
the experimentsand yield increasesin another28%of them], other workers could measureno effectsat all (Briggset al., 1926; Collins et al., 1929).
For 18 years Lemstrom (1904) studied the effects of exposure to electric
fields and reportedhealthy plants and increasedyields (includingcereals,
vegetablesand root crops). The time of day when electricitywas applied
was importantsince exposureat midday or in hot sunshinecould decrease
crop yields. However, a report of a subcommittee of the Board of Agricultureand Fisheries (Committee of Electroculture,Final reportof work
carriedout 1918-1937) indicated that, after prolongedresearch,benefits
of electricaltreatmentscould not be confidentlypredicted.
Recently Kruegeret al. (1978) have shown that the density and type of
air ions are important for plant growth, and suggestedthat ions rather
than electric fields producedthe various effects. Growth and respiration
rates (Kotaka et al., 1965) as well as ATP metabolism (Kotaka et al.,

187
1968) can be increased by air ions. Barley seedlings grown in ion-depleted

atmospheres (Krueger et al., 1965) show retarded growth, and plants
grown in Faraday cages (Hicks, 1957) are also weaker than ones grown
outside. Murr (1 963a, 1964a) studied the effects of large electric fields on
leaves of orchard grass (Dactylis glomerata) and reported burning of leaf
tips. He proposed that an over-respiration process was causing the burning, but Hart and Schottenfeld (1979) believed that the effects were due
to loss of water from the leaf tips when corona current flowed caused by
the high electric fields. Currents below 10-16 A had no effects on plants
(Murr, 1966a) while 10-15 to 10-9 A stimulated growth. Higher current
values destroyed the plants.
III.D (ii) Practical weed controlby high electricfields

Although plants have been killed by the application of very high electric
fields it is not possible to use this method for weed control. This method
requires arrays of wires suspended some 2 m or more above crops, all
charged to tens of kV. Outside of a laboratory this system is cumbersome
and dangerous. In addition broad-leaved weeds in cereal crops would be
affected less than the long, thin crop plants with pointed tips which would
be destroyed first.
If it can be shown that air ions stimulate growth then it may be possible
to use them, especially in greenhouses.
IV.
Microwaves and Weed Control
IV.A USES OF MICROWAVES IN AGRICULTURE
Microwave radiation has been suggested as a possible solution to many

and varied problems in agriculture, including prevention of frost damage,
rapid crop drying, reducing hard seed (impermeable seed coats), pest
control and weed control. Its advantages include the rapid penetration to
all parts of the "load," it leaves no residue after application and it can
be directed at its target. In weed control, microwave radiation is not
affected by winds, thus extending the periods of application compared to
conventional spraying methods. Furthermore, it can kill the roots of weeds
and also seeds buried to a depth of several centimeters in the soil as well
as nematodes and fungi. Although the primary concern of this review is
weed control some examples of other uses of microwave radiation are
indicated below, illustrative of the versatility of the technique.
In studies of the possible microwave protection of plants from cold,
Bosisio and Barthakur (1969) placed two wax bean plants (Phaseolus
vulgaris) in a cold chamber maintained at - 5?C for 4 hours. One plant
was treated with 2450 MHz radiation of power density 15 mW cm-2

188
THE BOTANICALREVIEW
which was sufficientto maintainthe leaf temperatureat 25?C.The authors

calculatedthat a power level of 2 mW cm-2 was being absorbedby the
leaf, and a minimum of 1 mW cm-2 representedthe threshold for leaf
protectionfrom frost at - 5?C.After transferto normal temperatures,the
irradiatedplant remained healthy whereasthe unprotectedone had been
destroyedby the cold. The investigatorsestimatedthat for frostprotection
125 kW of power would be needed per hectarewith high capital cost but
low runningcost.
In a later field test Bosisio et al. (1970) irradiatedfour-month-oldcorn
(Zea mays)in a plot 8 x 8 m and protected50%of it for 60 hours against
winds of between 8 and 33 km h- 1, temperatures of -1C
to
6?C and
1.5 cm of snow. A 2.4 kW generatorwas used and the irradiatingantenna

was erected 2 m above one corner of the plot. Power levels of 10 mW
cm-2 at 6 m from the antenna were the threshold for protection of the
crop. Plantsdirectlyexposedto the prevailingwinds were,however,killed,
even with 150 mW cm-2 radiation intensity.
Microwavetreatmentwould be very expensiveand pose healthhazards.
Although Bosisio et al. (1970) measured only 1 mW cm-2 at 3 m from
the edge of the plot, they had suggestedearlier (Bosisio and Barthakur,
1969) that farming areas using microwave radiation would have to be
kept clear of people during irradiations.Distributingthe energy equally
over large areas would also be a problem.
Boulangeret al. (1969) produced a design study of a microwave grain
drying system. Reducing grain moisture content from 20% to 15%took
less than 15 minutes by microwave methods (12 kW, 2450 MHz), 30
minutes by a high frequency(HF) dielectric system (13 MHz at 10 kW)
and 150 minutes by conventional hot-air techniques.Some wheat insects
and their larvae were also controlled (Tribolium confusum, Sitophilus
granarius and Cryptolestesferrugineus).Costs for the electronic system
were estimated to be half those of conventional methods. Baking and
milling tests showed little differencesbetween the methods. Fanslow and
Saul (1971), however, using a cavity with 1.8 kW at 2450 MHz and 0.6
kW at 915 MHz, found that there was a practicallimit to the increasein
drying speed since too rapid heating led to cracking of the kernel and
swellingof the grain,with consequentreductionin marketgrade.At very
high rates of dryingthey found that varyingthe air flow throughthe grain
had little effecton dryingrateswhereasBoulangeret al. (1969) reportthat
the dryingof wheatdependsupon the changein vaporpressuredetermined
by the air flow.
Microwave radiation has been used to kill insect pests. Hightower et
al. (1974) employed 2450 MHz radiationto control powder post beetles
in imported hardwoods,particularlyyellow pine and fir. Heatingrates of
30?Cmin'-I were achieved and the beetles and larvae could be controlled

ELECTROSTATIC
189
when wood temperaturesof 50?Cwere obtained. They predicted that a

30 kW unit could treat a 7.5 cm wide board at the rate of 1.5 m min-I
with costs comparableto kiln dryingmethods but considerablyfasterand
in less space. Nelson (1976a) and others (Iritani and Woodbury, 1954;
Nelson and Stetson, 1974; Nelson et al., 1966; Rai et al., 1971, 1972)
have studied the effects of radiation in treatinginsect pests in grain and
found that, often, much lower frequenciesof radiationare more suitable
(e.g. 39 MHz) becausethe dielectricpropertiesof insects and grainsdiffer
more from one another at lower frequenciesthan at microwave frequencies and so selective killing is easier. In relation to these investigations,
measurementshave been made on the dielectricand electricalproperties
of grains, seeds, insects, larvae and soils (Nelson 1973a, 1973b; Nelson
and Charity, 1972).
The proportion of hard seed in alfalfa (Medicagosativa) has been reduced (Nelson, 1976b) to levels of 5-15% from 40-60% following exposure to 39 MHz and 2450 MHz radiation. The success was related to
seed moisturecontent,the greatestreductionin hardseed beingin samples
with the lowest moisture content. The optimum temperaturewas found
to be 75?C.The benefits of treatmentlasted for up to four years after the
application.The effect was believed to be associatedwith the increaseof
watersorptionby the seeds. In studies on the seeds of trees (Kashyapand
Lewis, 1974), germinationhas been found to increase by 15%to 30%in
white and red spruce (Picea glauca and P. rubens)after exposure to 500
W for 20 or 35 seconds, depending upon the type of applicator.
Thomas et al. ( 1979) used a microwave oven for the rapid determination
of the moisture content of leaves of tobacco (Nicotiana). Samples were
exposed in a cavity for up to 2 minutes to 500 W of microwave power
at 2450 MHz. The results from microwave treatment were close to those
obtained by the Karl Fischer titrametric method but both of these nonthermal procedures gave somewhat lower values than those obtained using
a conventional hot oven. Temperatures in usual thermal techniques for
drying the samples (80-115?C) have, however, the disadvantage of releasing volatile non-aqueous substances as well as water from the leaf
tissue. Nicotine and propylene glycol in the microwave oven volatiles
were reduced by an order of magnitude compared to the amounts released
by conventional heating.
Diprose et al. (1979) dried leaf samples of Beta vulgaris and other
species in a microwave oven (2450 MHz, nominal 1 kW). Values for
moisture content fairly close to those from conventional measurements
were obtained after irradiation for 10 minutes. In Beta vulgaristhe leaf
temperaturewas found to rise from 200Cto nearly80?Cwithin 2 minutes

and then decrease to about 30?C after 5 minutes. It is suggested that the
relatively low temperatures may be beneficial in reducing the losses of

190
Table II
The effect of microwave radiation on seeds of four species (values give germination
percentage or seedling survival; from Davis et al., 1971)
4h
imbibed seed
Dry seed
Species
46 h
germinatingseed
45
270
23
45
23
45
J g-'
J g-l
J g-l
J g-l
J g-'
J g-I
0
Arachis hypogaea
100
83
53
46
Cucumis sativus
93
71
74
69
47
31
75
35
74
48
12
0
0
Echinochloa colonum
Amaranthus sp.
non-aqueous volatile substances. Other authors (Hankin and Sawhney,

1978; Miller et al., 1974; Routledge and Sabey, 1976; Steele, 1976) have
also reported the rapid drying of soils and foods (including determination
of moisture content) by microwave radiation.
Further applications include inhibiting the growth of microorganisms
and bacteria in flour (Olson, 1965) and the potential inhibition of mold
development on peanuts (Person, 1972). Jervis et al. (1974) have investigated the detection and control of grain loss in straw during combine
harvesting with microwave scattering techniques. They report that these
techniques are not suitable as the basis of a grain loss monitor mainly
because there are only small differences in scattered energy between grain
and straw.
IV.B
LABORATORY
EXPERIMENTS
WITH MICROWAVE
RADIATION
ON PLANTS AND SEEDS
Davis et al. (1971) irradiated seeds of twelve species (Zea mays, Arachis
hypogaea,Prosopisjuliflora, Cucumissativus, Brassica sp., Rumex crispus, Echinochloa colonum, Amaranthussp., Gossypiumhirsutum, Glycine max, Sorghum vulgare,Triticumvulgare)in a microwave chamber.
The magnetron source operated at 2450 MHz and 600 W. Samples of
dry seed, wet seed (4 hour imbibed) and germinating seed (46 hours) of
each species were investigated. Irradiation energies were measured with
respect to a 50 ml water load which gained 270 J g-I during a 60 second
exposure in the cavity.
There was a wide variety of responses but, in general, samples of dry
seed were less susceptible than 4 hour imbibed or 46 hour germinating
seeds (Table II). Dry seed could withstand six times the treatment of
imbibed seeds, e.g. for Arachis hypogaea, dry seed treated at 270 J g-'

191
Table III
Germination percentages and seedling survival for two (of eight) species whose
resistance to microwave radiation was greater for germinating seeds than for 4
hour imbibed seeds (from Davis et al., 1971)
Dry seed
45
270
J g-
JJg-
Zea mays
73
Sorghum vulgare
88
15
54
Species
4h
imbibed seed
23
-1
26
22
46 h
germinatingseed
45
23
45
J g-
J g-I
J g-1
52
62
15
gave 83% germination; 4 hour imbibed seed at 45 J g- ' did not germinate
at all; 46 hour germinating seed at 45 J g- I showed no seedling survival.
The other eight species showed similar results except that the 46 hour
germinating seeds were less susceptible than 4 hour imbibed ones (Table
III).
Fourteen-day-old plants of nine of the species were irradiated, damage
ranging from 3% of tissue (Brassica sp.) to 100% (Gossypium hirsutum)
for 9 J g-1 treatment. At 36 J g-' tissue damage was 25% and 100%
respectively.
Experiments were also carried out with combinations of dry and imbibed seeds in wet and dry sand. The condition of the soil containing the
seeds was found to be less important than the moisture content of the
seeds themselves in determining the susceptibility.
Davis et al. (1973) concluded that for 15 taxa [Canavalia ensiformis,
Phaseolus linensis vars. Hendersonbush and Fordhook242, Stizolobium

deeringianum and the twelve mentioned by Davis et al. (1971) excluding
Arachis hypogaea] seed susceptibility to microwave radiation was correlated with mass per seed, volume per seed, moisture content, ethersoluble content, and energy absorbed by the seed. The samples were
exposed to 600 W at 2450 MHz in a cavity and energy densities
were
again given in terms of energy gained by an equivalent water load. The

use of the energy density in terms of J g-I by Davis et al. (1971, 1973)
is, however, misleading. The figures given [23, 45 and 135 J g-' (Davis
et al., 1971)], do not represent the energies absorbed by the seeds. The
standard is reported as a 50 ml water load gaining 270 J g-I in 60 seconds.
Presumably the various energy densities were obtained by exposing the
samples for proportions of 60 seconds, e.g. 23 J g-' = 5 seconds, 135 J
g-
= 30 seconds.
The energy gained by a load depends upon many factors, among them

THE BOTANICALREVIEW
192
Table IV
The rise in temperatureof seeds exposed to 2450 MHz radiation.Results from

Davis et al. (1973, Fig. 2) but the AvTvalues are calculatedfrom the data and
equation (1); specificheat rangetaken between 1 and 2 J g-1 0C-1
Mass/
seed
Energy/
seed
(g)
(J)
Canavaliaensiformis
-1.5
-5.7
Zea mays
Prosopis juliflora
Triticum vulgare
-0.3
-4.2
14
--0.06
-2.2
-0.03
-1.7
Species
AT calc. ?C
s= I
3.8
s= 2
1.9
Germination
(%)
--4
-15
37
19
--49
57
29
--81
size, shape, moisture content, dielectric properties, mass of material being

exposed, and the frequency, strength and homogeneity of the irradiating
field (Nelson, 1973b; Olsen, 1975; Schrader and McNelis, 1975; White,
1970). If a water load absorbed a certain quantity of energy it does not
necessarily mean that another load in the same cavity for the same or a
different time would have absorbed the same or a proportional amount.
Calorimetry is a better method of determining energy absorbed by seeds
and exposure time in the cavity is another way of indicating the amount
of treatment.
Common to both of these studies (Davis et al., 1971, 1973) are dry
seeds of 11 species that were irradiated for 60 seconds (270 J g- 1). A
comparison of the germination results shows that for nine of them the
findings are very close but for two species they are different: Triticum
vulgare was reported to show 48% germination (1971) and 81% (1973);
Amaranthus sp. 35% (1971) and 99% (1973). Values for 4 hour imbibed
seeds of two species can be compared: Zea mays, exposure time 5 seconds,
germination 26%, and 10 seconds, 1%(1971) and 7 seconds, 22% (1973);
and Prosopis juliflora 5 seconds, I1%and 10 seconds, 1% (1971) and 7
seconds, 77% (1973). These values illustrate the variability that can arise
from microwave treatment. For the examples given the germination results for 1973 are higher than those expected from the 1971 ones. Such
differences can arise from variations in the operating conditions of the
microwave cavity and variations in seed batches, varieties or moisture
contents.
Davis et al. (1973, Fig. 2) plot germination against mass per seed and
energy absorbed per seed. No indication is given, however, of how the
latter was derived. Data taken from the Figure involved and used in the
equation:
E=m
x s x AT

(1)
193
Table V
Energy required for median toxicity in dried and imbibed seeds (after Rice and
Putnam, 1977)
Energydensity for LD50(J cm-2)
Species
Dry seed
Imbibedseed
Medicago lupulina
Portulaca oleracea
Echinochloa crus-galli
183
139
160
161
80
117
where E = energygained per seed (J)

m = mass of seed (g)
s = specific heat (J g-I 0C-1)
AT = temperature rise ?C
lead to the result that seeds with the lowest germinationhave the lowest
temperaturerises. Table IV gives some examples.
Lowestgerminationvalues would, however, be expectedfrom the seeds
with the highest temperaturerises. The discrepancycould be explained
if Davis et al. (1973) calculatedthe energyvalues per seed from exposure
times in the cavity and the energygained by the waterload in 60 seconds
ratherthan by measurement(e.g. calorimetricmethods).
Rice and Putnam(1977) investigatedthe effectsof microwaveradiation
on seeds of six species (Amaranthusretroflexus,Portulacaoleracea,Medicago lupulina,Digitaria sanguinalis, Echinochloa crus-galliand Setaria
italica). Samples were in a quartz holder placed in the waveguide and
microwave radiation was at a frequencyof 2450 MHz, the power level
being variable from 0.1 to 1.5 kW. Energyabsorption by the seeds was
determined by calorimetry immediately after irradiation. Values for a
50% decrease in germination (LDso) ranged between 88 J cm-2 (Digitaria
sanguinalis)and 183 J cm-2 (Medicago lupulina).Seeds of three species
were imbibed for 24 hours before irradiation.Median toxicity was then

reachedat lower energy levels (Table V).
In furtherstudiesseedsweremixed with threetypesof soil. The response
depended upon species, soil type, temperatureand soil moisture content
but, in general,the moister the soil the shorterthe treatmenttime required
to decrease germination. In addition, if the power of the radiation and
the exposuretime were varied to keep a constant power-time product,it
was found that the higher power (1.5 kW) was much more effective than
lower values (0.2 kW) for longertimes. With the sharpbend in the waveguide and the waterload placed as shown in Figure2 of Rice and Putnam
(1977), therewas likely to be some powerreflectedfrom these discontinui-

194
ties setting up a standing wave pattern in the waveguide. The introduction

of the sample holder into the region of maximum electric field would also
cause some power to be reflected. With standing waves present in the
waveguide, the position of the sample holder is important in determining
the amount of power dissipated in the sample arising from the variations
in electric field intensity that occur. Since Rice and Putnam report no
reflected power it must be presumed that suitable tuning was used to
minimize the reflections and even out the power flow.
The method of measuring energy absorbed by the load (i.e. calorimetry)
is preferable to stating exposure times or comparisons with a standard
water load, especially when the samples are placed in a waveguide for
irradiation. The energy values measured by calorimetry would be Joules
or Joules g-'. The values given in Table 1 of Rice and Putnam are in
Joules cm-2 but no indication is given of how these values are derived
or to what particular area reference is being made.
Bhartia et al. (1977) exposed various grain and oil seeds, including
Torch rape (Brassica napus), Norland flax (Linum usitatissimum), and
wild oat (Avena fatua), to 360 seconds of 2450 MHz radiation. Torch
rape was little affected by the treatment. In Norland flax and wild oat,
however, germination was reduced to 15% and 5% respectively 96 hours
after irradiation compared to 88% and 50% respectively for the controls.
The authors suggest that irradiation of seeds prior to sowing could give
effective weed control.
Diprose et al. (1978a) suggested that wild oat (Avena fatua) might be
controlled selectively in cereal crops. They reported a decrease in germination of Avena fatua from 60% to 30% when exposed to 30 seconds
of 2450 MHz radiation (germinated in Levington compost) and from 20%
to 15% when similarly exposed and germinated on moist filter paper in
petri dishes at 25?C. The power level to the cavity was a nominal 500 W,
sufficient to dissipate 5 J cm-3 s-I in a 40 ml water load. Further work,
however, with more replications and Avena fatua from different sources
and harvested in different years showed the results were not consistent
(unpublished observations). The germination of various samples of irradiated grains of wild oat varied, ranging from 0% to 60%. Low germination values were then attributed to increased dormancy rather than
increased mortality.
Lal and Reed (1980) exposed seeds of wild oat (Avenafatua var. montana) to power levels of 0.1 to 1 kW of 2450 MHz microwave radiation.
The seeds, in a glass sample holder, were placed in a length of waveguide
terminated by a waterload. A maximum of 50% of the seeds was destroyed
after 30 seconds of exposure to 1 kW (26.3% moisture content). A similar
exposure time resulted in death of 38% and 14% of wild oat seeds of
18.9% and 7.4% moisture contents respectively. Seeds kept in soil for 72

ELECTROSTATIC
195
Table VI
The percentageof seeds destroyedafterexposureto 1 kW of 2450 MHz radiation
for 10, 20 or 30 seconds (afterLal and Reed, 1980)
Seed
moisture
content
% seed destroyed
(%)
10 s
20 s
30 s
Harman oats
17.2
7.5
63
5
80
33
90
50
Cypress wheat
17.8
44
77
3
7
24
Wild oats
6.5
18.9
12
88
58
40
7.4
13
Seed type
hours at 1?Cprior to exposure, and tested when still within the soil, were
more susceptible: 100%, 95% and 90% of the seeds being killed in soil at
20.2%, 17.3% and 13.8% moisture contents respectively.
Other experiments showed that Harman oats and Cypress wheat were
far more susceptible than wild oats to 1 kW of 2450 MHz radiation at
both low and high moisture contents (Table VI).
Bigu-del-Blanco et al. (1977) exposed seedlings of Zea mays (var. Golden Bantam) to 9000 MHz radiation. Forty-eight-hour old seedlings in
plastic bags were treated with microwave radiation for 22-24 hours. The
exposure chamber was a microwave anechoic chamber and power density
levels were between 10 and 30 mW cm-2 at the point of exposure in the
absence of the seedlings. Anechoic chamber temperature and humidity
were controlled. Temperature increases up to 4?Cwere measured in treated
specimens when compared with control seedlings in the chamber. After
treatment, irradiated specimens appeared dehydrated and with a rugose
texture. Control seedlings remained healthy and showed about one-fifth
of the moisture loss of the treated seedlings. Not only were the seedlings
different in appearance but when planted in perlite there was a difference
in growth (irradiated retarded), but this disappeared after a few weeks.
The authors concluded that the long exposure to 9 GHz microwave radiation, even at low powers, was sufficient to dehydrate the seedlings so
inhibiting their development.
Crawford (1977) examined the phytotoxic effect of 2450 MHz radiation
on seeds of Trifolium and Medicago (three cultivars of Trifolium repens
and cultivar Eynsford of Medicago sativa). The seeds were exposed in a
microwave oven with a nominal 1.2 kW input; a 100 ml water load had
690 W dissipated in it at the power setting used. Treatment times were
between 5 and 45 seconds. All samples showed a marked decrease in

196
THE BOTANICALREVIEW
germination after treatment for 15-20 seconds. Numbers of hard seeds

decreasedfor exposuretimes of between 5 and 15 seconds.Aftertreatment
for 45 seconds, germinationof the seeds of all the cultivarswas very low,
none exceeding 15%.
IV.C FIELD EXPERIMENTS WITH MICROWAVES FOR WEED CONTROL
Waylandet al. (1973a) planted seeds of wheat (Triticumaestivum)and

radish (Raphanussativus)in packets buried 2.5 cm in Lufkin fine sandy
loam soil of 6.8% moisture content. The seeds had been imbibed for 10
hours prior to burial. Microwaves were applied to the samples from a
1500 W (maximum) source at 2450 MHz via a 10.2 cm square, 5 cm
deep, stainless-steelradiator.The pre-treatmentsoil temperaturewas 25?C.
It was found that 100 J cm-2 (Triticumaestivum)and 180 J cm-2 (Raphanus sativus)were needed to produce 50%mortality in the seeds. At 210
J cm-2, germinationvalues were reducedto 15%and 40%for wheat and
radish respectively.
In separateexperimentsconstanttreatmentsof 210 J cm-2 were applied
to samples of seeds but at differentpower levels. For wheat it made no
appreciabledifferencewhether 100 W was used for a long time or 1200
W for a short one: the percentageof seeds killed was approximately60%
in each case. In radish,however, 35%of seeds werekilled at 100 W, rising
to 45% at the 1200 W power level. Soil temperaturesvaried from 60?C
at 100 W to 50?Cat 1200 W after the constant 210 J cm-2 treatments.
The extent of destructionof radish seeds was not the same in the two sets
of experiments.At 210 J cm-2 about 60% were killed in the first experiments and approximately40% in the second. For wheat, however, the
values varied from about 15%survival in the first to more than 40% in
the constant energy,differentpower experiments.
Waylandet al. (1975) continued their field experimentswith four, 1.5
kW, 2450 MHz generatorsmounted in a four-wheeled1.2 x 2.4 m trailer.
The outputswereconnectedto a 20.4 x 20.4 x 5 cm radiator,the opening
of which was covered by a ceramic sheet. Experimentswere carriedout
at three field sites in Florida and Texas [Freeport(Lakelandsand, 10%
moisture content, 23?Csoil temperature),Weslaco (Hidalgo sandy loam,
12%,25?C)and College Station (Lufkinsandy loam, 20%, 20?C)].
Seeds of Triticum aestivum (wheat), Hordeum vulgare (barley),Avena
byzantina (oats), Brassica hirta (mustard), Raphanus sativus (radish) and
Brassica napus (turnip)were sown with the addition of Zea mays (corn)
and Sorghum bicolor(sorghum).The microwave generatorswere drawn
over the plots to give treatmentsof 35 to 325 J cm-2. To obtain consistent
pre-emergencecontrol (>80%) of both broad-leavedplants and grasses
at least 183 J cm-2 was needed.

197
Post-emergent tests were carried out at Weslaco. Young plants of Echinochloafrumentacea(Japanese millet), Sisymbriumirio(London rocket),
Helianthus annuus (sunflower), Euphorbiaglyptosperma(ridgeseed euphorbia) and Amaranthus retroflexus(redroot pigweed) were approximately 14.3, 2.6, 4.8, 0.3 and 4.6 cm high respectively at the time of
treatment (27 days after planting). Energy levels of 77, 154 and 309 J
cm-2 were applied at a power level of 4.4 kW. Eight days after irradiation,
treatment with 77 J cm-2 had completely controlled sunflower, ridgeseed
euphorbia and redroot pigweed. Japanese millet was controlled to the
extent of 73% and London rocket 87%. The two last species were controlled by over 90% with 154 J cm-2 and completely by 309 J cm-2. The
unactivated radiator, however, caused 12%injury to the plants as it passed
over them.
At Freeport there was a natural stand of plants of "cut-leaved primrose"
(Oenotheralaciniata) 5-15 cm high; when treatments of 70 J cm-2 were
applied, 99% of them were killed. The same treatment to young wheat
plants 10 cm tall killed 73% of them. Established "primroses" were more
susceptible than young wheat. A similar trend was observed in glasshouse
studies with morning glory (Ipomoea sp.), redroot pigweed, Texas panicum (Panicumtexanum)and barnyard grass (Echinochloacrus-galli);15day-old plants were more susceptible than 8-day-old seedlings, the greater
susceptibility probably being associated with the larger leaf surface area.
Experiments with Johnson grass (Sorghum halepense) in Miller clay
soil showed 100% control of top growth with 560 J cm-2. After two weeks
there were 48 new seedlings im-2 and 11.5 sprouts from rhizomes m-2.
Energy levels of 2240 J cm-2 also resulted in 100% top kill but left only
1.5 seedlings mi-2 and no rhizome sprouts 14 days afterwards.
Table VII shows that there are considerable variations from place to
place in the response of buried seeds to microwave radiation. At the 70
J cm-2 energy level and 4 kW power level, for instance, control of broadleaved weeds ranges from 21-75% and of grasses from 8-60%. At higher
power levels, e.g. 183 J cm-2 (4 kW), control becomes more consistent
(between 80 and 91%) for both types of seed. It is evident that grasses
that have started development are more susceptible to 70 J cm-2 than
buried seeds. At Freeport 10-day-old wheat seedlings were controlled to
the extent of 73% (4 kW) and 67% (5.2 kW) compared with the poor
control of sown wheat grains of 8% (4 kW) and 24% (2 kW). Established
"primrose" was controlled to the extent of 99% (4 kW) compared with
35% for the seeds of other broad-leaved plants.
Plants established above ground are the easiest to kill, as with hoeing,
flaming or spraying. The advantage of microwaves (i.e. penetration into
the soil) is offset by the amount of energy required.
Menges and Wayland (1974) compared weed control by microwaves

THE BOTANICALREVIEW
198
Table VII
Percentage control of plants treated with different energy and power levels (data
from Wayland et al., 1975)
Control (%)
Power
Energy level
(J cm-2) (kW)
70
2
4
Broad-leaved weeds
Freeport
Weslaco
College
Station
Freeport
75
0
21
35
24
8
75
4
4
144
5.2
4
183
325
5.2
94
-
71
-
91
-
80
-
Wes- College
laco Station
30
51
60
"Primrose" 99*
Wheat 73*
Wheat 67*
71
24
77
60
37
89
87
81
-
91
-
60
88
83
5.2
141
Grasses
Results are for treatment of seeds in soil except for those marked * which were treated
after emergence.
Soil moisture contents and type: Freeport = 10%-Lakeland sand, Weslaco = 12%-Hidalgo sandy loam, College Station = 20%-Lufkin sandy loam.
with control by chemical herbicides in vegetable crops. Smooth flat beds

were prepared in Hidalgo sandy loam soil and various herbicides were
sprayed on the beds and then incorporated 3.8 cm into the soil. Seeds of
London rocket (Sisymbrium irio) and sunflower (Helianthus annuus) were
buried 0.3 cm and 1 cm deep respectively after the herbicide treatments.
Other weed-seeded plots were treated with microwave radiation at 2450
MHz at rates from 45 J cm-2 (electronic carriage speed 0.0384 km h-')
to 720 J cm-2 (0.0024 km h-'). Four hours after ultra high frequency
(UHF) treatment "Yellow granex PRR" onions were seeded in all the
plots. Of the herbicides only methazole (2.2 kg ha- 1) controlled London
rocket (99% killed) but it also damaged the onion (85% damage) and only
72% of the sunflower were killed. The next most effective herbicide for
London rocket was a propachlor and mulch treatment (9.0 kg ha- 1)which
killed 48% of the London rocket and 20% of the sunflower with 30%
damage to onions. Sunflower was most successfully controlled by methazole and next best was perfluidone (3.4 kg ha-') killing 36% of the
sunflower but with 85% injury to the onions. Microwaves at 360 J cm-2
killed 94% of the London rocket and all of the sunflower. Damage to the
onions was 18% which compared with 12% and 17% damage to them in
the weeded and non-weeded control plots.
Sisymbrium irio, Euphorbia glyptosperma, Amaranthus retroflexus,
Echinochloa frumentacea and Helianthus annuus were seeded in both

199
irrigated and non-irrigated plots and irradiated with 2450 MHz microwaves. Cantaloupe (Cucumis melo var. reticulatus) was seeded 2 cm deep
one day after UHF treatment. In the irrigated plots 45 J cm-2 gave
inadequate weed control but 90 J cm-2 produced over 80% control of all
the weeds except for common purslane (Portulaca oleracea), a volunteer
which was controlled to the extent of 50%. One hundred and eighty J
cm-2 produced greater than 96% control except for common purslane at
81%. The soil temperature was 80?C 1 minute after irradiation and 44?C
10 minutes after exposure (180 J cm-2). The cantaloupe size and stand
were visually estimated to be greater (after treatment at 90 and 180 J
cm-2) than the control.
In the non-irrigated soil all weed species were controlled by 360 J cm-2
and 720 J cm-2 and all except common purslane (77%) at 180 J cm-2.
Soil temperatures reached 103?C (3.8 cm deep) 1 minute after 720 J cm-2
treatment and 47?C after 150 minutes. Cantaloupe yields were increased
in the 180 (150%), 360 (120%) and 720 (140%) J cm-2 plots as compared
with non-weeded control plots.
Menges and Wayland (1974) broadcast-seeded redroot pigweed (Amaranthus retroflexus), and disked and bedded them up to 13 cm deep in
the soil (sandy loam). London rocket (Sisymbrium irio) and sunflower
(Helianthus annuus) were seeded 2.5 cm deep into irrigated plots. The
irrigated plots were watered daily until microwave treatment, after which
"yellow granex" onions were planted. Seeds in irrigated soils were more
susceptible to the microwave treatment with both London rocket and
sunflower being controlled by 360 J cm-2 (approximately 90% mortality
for both in dry soil). With a treatment of 180 J cm-2, control of London
rocket in wet and dry soils was reduced to 87% and 66% and sunflower
to 93% and 68% respectively. The stage of seed imbibition was thought
to be more significant than soil moisture content.
Redroot pigweed was controlled by a treatment of 360 J cm-2 down
to 7.5 cm in both irrigated and non-irrigated soils and there was an effect
down to 10 cm in the dry soil. At 180 J cm-2 the radiation was effective
to 7.5 and 5.0 cm in the dry and wet soils respectively. At 90 J cm-2 the
radiation was not effective in the irrigated plots and seemed to increase
the germination in the non-irrigated soil. Onion yield was increased at
the 90, 180 and 360 J cm-2 treatment levels by 161%, 272% and 233%
respectively over the non-weeded "check," but, when compared with a
hand-weeded area, the three treatments gave 833%,137% and 120% yields.
Whatley et al. (1973) found that in Sawyer fine sandy loam a low
moisture content (<0.5%) attenuated microwaves less than the same soil
with 10-13% moisture present (2450 MHz). Whatley recommends that,
when used as a pre-emergence treatment, microwaves should be applied
when the top 1-2 cm of soil contains minimum moisture. In a later paper

200
Whatley et al. (1974) claimed that in laboratory experiments minimum

soil moisture and maximum seed moisture are ideal. In field applications
high seed moistures and soil moistures are desirable. Soil type is important
when oven-dried soils are used in microwave experiments, but in practice
when they are wet the differences are less important than the moisture
content of the seeds. Champ et al. (1972) used 2450 MHz radiation to
control aquatic weeds. Initially, duckweeds (Wolffia punctata) were exposed in a cavity to 600 W for 8 seconds (16.7 J cm-3) and half the plants
died. All the plants died after treatment for 10 seconds. In simulated field
conditions, control of Spirodela sp., Wolffiapunctata and W. columbiana
was demonstrated although no estimate of energy requirements was given
except that they would be much higher than in cavity experiments. The
plants died over a period of several days, a useful feature in aquatic weed
management, as rapid death and decay of plants can seriously disturb the
oxygen levels in the water leading to the death of fish (Price, 1976).
Hightower et al. (1974) used four different types of seeds (two grasses,
a clover and turnip) in experiments with 2450 MHz radiation. Samples
were spread on a soil surface and microwaves applied from a horn antenna
at power levels from 100 W to 750 W for periods between 2 and 20
minutes. These gave power density levels between 50 and 300 J cm-2.
There was little or no control of germination. The seed type most affected
was turnip whose germination stayed constant at 78% up to an energy
level of 200 J cm-2 but fell to 60% at 300 J cm-2.
In further experiments seeds of Kentucky tall fescue grass (Festuca
elatior) were placed on a soil surface in an area 5.2 x 5.3 cm which was
2 cm beneath a dielectrically loaded waveguide. Power densities could be
varied from 100 J cm-2 to 24,000 J cm-2. Three kinds of soil were usedFuquay, Greenville and Tifton-representing sandy, sandy loam and clay
types. Exposure times were from 20 to 60 seconds at a power level of 400
W. No appreciable reduction in germination was recorded for seeds on
any of the soils.
The power was increased to 500 W and irradiation for 6 minutes was
needed before the germination of seeds on Tifton soil began to drop. After
10 minutes it was reduced to 78%. Germination of seeds treated on
Greenville soil rose steadily from 88% (5 minutes) to 94% (10 minutes).
When 1000 W was applied, seeds on Fuquay and Tifton soils were not
noticeably affected. Seeds in Greenville sandy loam started to be killed
after 5 minutes (95% germination) but 72% still germinated after exposure
for 10 minutes.
Seeds imbibed for 12 hours were then laid on damp soils. Kentucky
fescue was completely controlled after 2 minutes (Greenville soil) at a
750 W power level and after 8 minutes on Tifton soil (90% germination

201
after treatment for 2 minutes). When 1200 W was used an irradiation

time of less than 1 minute gave very considerable control of seed germination on both Greenville and Tifton soil. Hightower et al. (1974)
calculatedthat control of damp seeds on damp soil needed 1500 J cm-2
at the 750 W level for Greenville soil and 10,500 J cm-2 for Tifton soil.
At 1200 W 1600 J cm-2 was needed for both soils. The authors stated
that at these rates a machine producing 2 kW of power would have to
travel at 0.00035 km h-1 to treat one acre of dry soil thus taking 8750
hours to complete the task. In general,however, the results of Hightower
et al. (1974) agree with those of previous authors:moist seeds are more
susceptible than dry ones. When wet, the difference in two soil types
(Greenville and Tifton) became much less markedthan when they were
dry, in agreementwith the conclusions of Whatley et al. (1974).
The irradiationtimes- 10 minutes at 1 kW for dry soils and seedsare long, but others have reportedsimilar times, especially for dry seeds.
Bhartia et al. (1977) treated seeds in a cavity for 360 seconds at 1 kW
and killed those of several species, but not all. Shaferand Smith (1974)
treated seeds of foxtail millet (Setaria italica) with 1 kW of 2450 MHz
radiation. Dry seeds had an LD50 of 30 minutes, and 4 and 16 hour
imbibed seeds had LD5_'sof 35 and 25 seconds respectively.
Davis et al. (1973) showed that small seeds are less susceptible than
large ones, so seeds of Kentucky fescue would be expected to be harder
to kill than, for example, wheat. Dry, small seeds are the hardest of all
to kill and so Hightoweret al. (1974) chose a difficultexample for their
experiments.Hightoweret al. mention treatmentsup to 24,000 J cm-2.
This seems high but is only 240 J mm-2. Also this is the energyincident
upon the seed in the area-not all of it is absorbedso the amount absorbed
by the seed could be very small.
The equipmentused by Hightoweret al. is differentfrom that employed
by Waylandet al. (1973a). The latterworkersused a small cavity (10.2 x
10.2 x 5 cm) placed on the soil surface.With this they claim that 200 J
cm-2 is adequate for general weed control and up to 600 J cm-2 for
difficultweeds such as Johnson grass (Sorghumhalepense).Hightoweret
al. claim that a minimum of 1600 J cm-2 is necessarywith their experimental apparatus. They used a length of dielectric loaded waveguide
separatedfrom the seeds by 2 cm. More information is necessary,however, such as moisture contents of soils and seeds, and temperaturesof
soils and seeds after irradiation,to allow any detailed comparison with
other results.
Menges and Wayland (1974) measured the soil temperatureafter irradiationwhen using a 1.5 kW generator.One minute after treatmentat
180 J cm-2 the soil temperaturewas 80?C.The effect would help to kill

202
any seeds in addition to the dielectric heating within them. By putting

seeds on the surface, Hightower et al. (1974) lost the benefit of the bulk
heating of the soil.
IV.D
THE EFFECT OF MICROWAVE
MICROORGANISMS
RADIATION
UPON SOIL
AND NEMATODES
Wainwright et al. (1980) exposed small (20 g) samples of various soils

(an organic loam, Fitzwilliam and Chapeltown brown earths) to 1 kW of
2450 MHz radiation in a cavity to measure the effect upon the microorganisms. As the exposure times were increased from 10 to 30 seconds
nitrification in organic loam soil was progressively reduced, with an accompanying rise in NH4+-H. Respiration rates in the organic loam and
Fitzwilliam brown earth were slightly depressed after a 20 second exposure
but not by a statistically significant amount. Sulphur oxidation was stimulated in Chapeltown brown earth after a 20 second exposure to 2450
MHz radiation and the number of S-ions in the organic loam increased
two- and three-fold compared with the control 28 days after treatment
for 20 seconds.
Plates prepared from the organic loam showed that fungi were reduced
from 30,000 g-I to zero after a 20 second exposure in the cavity while
the numbers of heterotrophic bacteria remained unaffected. Numbers of
thiobacilli increased considerably, treated soil containing up to nearly five
times as many as the control 28 days after treatment. It was concluded
that the microwaves had marked differential effects on the soil microorganisms, were unlikely to harm the soil, and might stimulate seedling
emergence and growth.
Diprose et al. (1978c) irradiated small samples (ca. 25 g) of organic
loam in a microwave oven cavity (46 x 30 x 34.5 cm) with a nominal
1 kW of 2450 MHz radiation (corresponding to 11 J cm-3 s-I dissipated
in a 40 ml water load in a 6 cm diameter petri dish). Fungal colonies
were eradicated at exposures of 30 seconds and above, while bacterial
colonies remained unaffected at 30 seconds, were reduced to ca. 20% of
control values after 120 seconds and were almost all killed (ca. 2% survival) at 360 seconds. Lyon et al. (1979) inoculated sterilized samples of
organic loam and John Innes compost (both at 25% and 50% moisture
contents) with spores of Botrytis cinerea and Fusarium. Samples were
placed in a microwave oven cavity and exposed for 4, 8 or 12 seconds
to a nominal 1 kW of 2450 MHz radiation. Spores in both soils at the
higher moisture content were more susceptible than when in soils at 25%
moisture. Fusarium appeared more resistant than Botrytis-in organic
loam after a 12 second exposure, Botrytis showed only 16% (25% moisture) and 0% (50% moisture) spore survival while 76% and 14% respec-

203
tively of the Fusariumspores were still active. Post-irradiationtreatment

influencedthe number of spores killed. Samples irradiatedfor 6 seconds
were allowed to cool in a refrigerator(5?C)and placed on a bench top
(20?C)or in a well insulatedbox. The numbersof fungalspores surviving
were 19.2%,20.9% and 5.3%respectively.
Vela et al. (1976) used differenttypes of soil (including sandy loam,
loam mix and clay) from severallocations in Texas to determinethe effect
of microwavesupon soil microorganisms.Samples(ca. 250 g) were placed
in a microwave cavity and exposed to 1 kW of 2450 MHz radiation for
5, 10, 20 and 40 seconds, and soil temperaturesreached 80?C.
From the results of all the soil types, only fungi showed a statistically
significantreductionwith increasingexposure.All other organisms(bacterialspores,nitrogen-fixingand otherbacteria,actinomycetes)were little
affectedregardlessof soil type or location.
Microorganismsin wet soils (at water holding capacity) were more
susceptiblethan in dry soils (3 to 7%moisturecontent).Even so, exposure
for at least 60 seconds was needed (60,000 J to the cavity) before large
reductions in populations occurred. Fungi were eliminated in both wet
and dry soils after 120 seconds and in wet soils after 60 seconds. After
480 seconds there were still microorganismsalive although 100 to 1000
times fewer than in the control (e.g. total bacterial count: control, dry
soil, 1.3 x 106 per g soil; 480 seconds, dry soil, 2.9 x 103per g soil). In
wet soils, however, short treatments(e.g. 10 or 20 seconds),increasedthe
activities of bacteria and fungi relative to those of dry soils. Although
bacteriacould withstandexposure for 8 minutes to 1 kW of microwaves
when in soil, isolated Azotobacterin aqueous cultureswere killed after a
20 second treatment.
Field studies were carried out with a Phytox CorporationZapper III
60 kW machine at several sites. Energy density treatments were 200, 400
and 800 J cm-2. These were all sufficient for good control of weed populations but the microwave radiation did not seriously disturb the soil's
autochthonous microflora. It may be noted that Vela et al. (1976) present
energydensities in thousandsof J cm-2, e.g. 60,000 J cm-2, for laboratory
experiments. However, the values should be just the number of Joules,
i.e. 60,000. This corresponds to 1 kW for 60 seconds (G. R. Vela, pers.
comm.). The figures given for the field trials are correct.
O'Bannon and Good (1971) placed small samples (90 cm3) of a composted glasshouse soil (1:2:3 composted manure: sandy clay loam: coarse
sand; 11% moisture content) infected with root knot nematode (Meloidogyne incognita) in a microwave cavity. Exposure to 1250 W of 2450
MHz radiation between 30 and 300 seconds eliminated the nematode
population. Exposure for 15 seconds severely reduced activity. Similar
results were obtained when larger (400 g) amounts of soil were used.

204
In studies by Heald et al. (1973), soil infested with Rotylenchulusreniformis (2300 R. reniformis100 g-') was placed in plastic bagswhich were
buried 2.5 and 8 cm deep in soil. The ground was then irradiatedwith
2450 MHz. Nematode counts showed a significant decrease at energy
densities of 732 and 322 J cm-2 but not at 183 J cm-2. Plants grown in
treatedplots were reportedto be largerthan those in non-treatedsoils.
The effect of microwaves at 2450 MHz was compared with that of a
chemical fumigant (1,3-dichloropropene)by Heald et al. (1974) in the
treatmentof nematodes in the ground.The soil was a Hidalgo fine sandy
loam naturally-infestedwith Rotylenchulusreniformisand common purslane (Portulacaoleracea).The fumigantwas applied 20-25 cm deep at a
rate of 74.8 1 ha-'. Microwaves from a 60 kW source were applied at
400 and 800 J cm-2 and one trialinvolved both fumigantand UHF (1200
J cm-2). Southernpeas ( Vignaunguiculatassp. unguiculatacv. Blackeye)
were planted after the treatment.
Microwavesat 400 J cm-2 gave 31%control of common purslane,but
did not affectthe yield of peas. Nematode counts 50 days aftertreatment
showed populations about eight times the size before treatmentbut this
fell to about four times after 82 days. Common purslane was almost
completely controlledby 800 J cm-2 but this level of treatmenthad little
overall effect on the nematodes. Yields and sizes of pea were, however,
increased.The fumigantalone was the most effectivein reducingnematode
infestations but had no effect on the weeds. Yield and size of pea were
increased as they were also when both UHF and fumigantwere used in
association. Very good control (99%)of common purslanewas achieved
but the counts of Rotylenchulusreniformiswere three times higher after
82 days than with the fumigant alone. It was suggestedthat fumigants
may be effective for nematode control at depths 10 to 25 cm below the
soil surfacein combination with microwaves used for control in the top
10 cm of soil.
IV. E CONCLUSIONS
IV.E (i) Summary

Davis et al. (1971, 1973) demonstratedthat for 15 species the susceptibility of seeds exposed in a cavity to 2450 MHz radiation depended
upon the mass, volume, moisture content, ether-soluble content of the
seeds and the energy absorbed by them. Other authors have similarly
described the phytotoxic effect of microwave radiation (Bhartia et al.,
1977; Crawford,1977; Diprose et al., 1978a; Rice and Putnam, 1977).
When seeds are in the soil they can still be killed by microwaves.When
the soil is dry its type is importantin determiningthe amount of radiation

205
needed to kill the seeds. The differences between soils are progressively
less marked as they become wetter or as the amount of energy provided
is increased (Rice and Putnam, 1977; Wayland et al., 1975; Whatley et
al., 1973). The state of imbibition of buried seeds affects their response
to microwaves. Moist seeds are more susceptible than dry ones whether
in wet or dry soils, and seeds in wet soils are more easily killed than in
dry ones. Large, imbibed seeds in wet soils are the easiest to kill and small
dry ones in or on top of dry soils the most difficult (Hightower et al.,
1974; Menges and Wayland, 1974).
The depth to which treatment is effective in soils obviously depends
upon how far the radiation can penetrate. The soil type, its moisture
content and distribution, density and compaction influence depths of
treatment as well as the frequency and strength of the radiation (Cihlar
and Ulaby, 1974).
Radiation is absorbed more rapidly by wet soils than dry ones. Menges
and Wayland (1974), using a treatment of 360 J cm-2, found it effective
down to 10 cm in dry soils and 7.5 cm in irrigated ones. At 180 J cm-2
penetration was down to 7.5 and 5.0 cm in dry and wet soils respectively.
For a given soil condition the only way to increase the depth of treatment
is to increase the energy density. In a practical situation this means that
any device would have to monitor soil conditions carefully as it progressed, otherwise the effect could vary since it is not uncommon for soils
to differ considerably between one part of a large field and another. In
addition, a device would desirably be operated to get the maximum effect
for a minimal expenditure of energy and, since soil conditions are so
important, constant monitoring would be necessary to ensure effective
treatment. Under the varying soil conditions, high powers and large energy
densities would have to be used to make sure all seeds are killed. The
advantage of microwave weed control systems over flaming or hoeing is
its penetration into the soil but this is offset by its dependence on soil
conditions and the large amounts of energy required.
Plants can be killed by microwaves (Champ et al., 1972; Davis et al.,
1971; Wayland et al., 1972). Broad-leaved weeds are more susceptible
than grasses, especially at lower energy levels, but the difference is not
enough to lead to a useful selective method of control (Wayland et al.,
1975). The greater susceptibility of broad-leaved species may result from
the greater absorption of energy by their larger surface area in comparison
with those of grasses for example.
Microwave radiation has been used to control nematodes (Heald et al.,
1974; O'Bannon and Good, 1971) but it is not as effective as fumigants.
In addition, if only strips are irradiated, re-infestation will take place from
neighboring untreated soil. Fungi have also been successfully controlled

206
(Diprose et al., 1978c; Lyon et al., 1979; Vela et al., 1976) without killing
biologically important bacteria and little damage is done to the soil (Wainwright et al., 1980).
IV.E (ii) Thermal or non-thermal mechanisms of death
The precise cause of death of seeds and plants cannot be definitely
stated. Both thermal and non-thermal mechanisms have been proposed.
Wayland et al. (1972) exposed imbibed seeds of "broadleaf bean" (Phaseolus vulgaris var. Black Valentine) and leaves of three-month-old mesquite (Prosopis glandulosa) to 600 W of 2450 MHz radiation. The seeds
rose in temperature to 50?C and only 38% germinated; however, the
germination of similar seeds in a water bath at 50?C was not affected.
Similarly, leaves of mesquite heated to 46?C by microwaves were damaged, but leaves in an oven at 46?C showed no tissue damage after exposure
for 5 minutes. The authors concluded that selective heating or non-thermal effects occurred.
In further experiments Wayland et al. (1 973b) exposed samples of beet
(Beta vulgaris) to 600 W at 2450 MHz and immersed other samples in
water baths. Cell membranes were disrupted but whether by thermal or
non-thermal means was not clearly distinguishable, although membrane
disruption by microwaves apparently occurred at temperatures 20?C below similar disruption in a water bath.
Schrader and Patel (1977) investigated the loss of viability of peas
(Pisum sativum) when exposed to 916 MHz radiation up to 20 kW power
levels. Comparing the results with those obtained by heating peas in
conventional high temperature ovens, they concluded that there was little
likelihood of a significant non-thermal effect. Diprose et al. (1978a) irradiated 100 grains of wheat (Triticum aestivum) with 1 kW, 2450 MHz
radiation for 180 seconds and another 100 grains for six periods of 30
seconds each with 15 minute intervals between exposures. The continuous
exposure killed 84% of the grains compared with 25% of those periodically
exposed. When the experiment was repeated with grains of wheat, barley
and oats, but with a one hour interval between each irradiation, germination values for the continuous exposures were 6%, 4% and 2% and for
the intermittent ones 92%, 96% and 72% respectively. The authors believed that these results indicated principally thermal effects on the grains.
Goldblith and Wang (1 9 6 7) compared the effect of conventional heating
and of microwaves (2450 MHz) on the inactivation of Escherichia coli
and spores of Bacillus subtilis, and found essentially the same results. In
line with these observations, Lechowich et al. (1969) found similar population decreases of suspensions of microorganisms (Streptococcus faecalis and Saccharomyces cerevisiae) when irradiated by microwaves and

207
when heated conventionally. Vela and Wu (1979) also concluded that soil
microorganisms were killed only by thermal means when heated by a
nominal 1 kW of 2450 MHz radiation. Olson (1965), however, stated
that growth of bread molds was inhibited by microwaves and the effects
were probably not due to conventional thermal means. Baker and Fuller
(1969) experimented on the control of soil-borne pathogenic fungi with
2450 MHz radiation but were uncertain as to whether there was a nonthermal effect in addition to a thermal one.
Cope (1976) suggested that the evidence of superconductive properties
of biologically important organic substances at high temperatures and the
microwave effects on superconductive tunnelling in carbon films present
the possibility of non-thermal effects of microwaves on biological systems.
Mayers and Habeshaw (1973) observed the depression of phagocytic activity when cultures of mouse macrophages perfused with suspensions of
human red blood cells were exposed to 50 mW cm-2 of 2450 MHz
radiation. The activity returned to normal when the radiation was discontinued, but the depression could not be accounted for thermally, since
the 2.5?C rise in temperature found during irradiation would not have
been expected to increase the activity. Sher et al. (1970) compared the
effects of Joule heating and field-induced forces on "pearl-chain formations" of microscopic particles. They concluded that Joule heating would
predominate over field-induced effects in biological structures smaller
than 50 ,um, excluding the possibility of non-thermal genetic damage by
field-induced force effects from microwaves.
Whether thermal, non-thermal effects, or a mixture of these are the
cause of death of seeds and plants when irradiated by microwaves is not
clearly resolved. When practical microwave weed killers have been used
soil temperatures were found to be quite high-80?C or more (Menges
and Wayland, 1974). High temperatures in seeds have been reported when
irradiated in cavities (Diprose et al., 1978a; Schafer and Smith, 1974).
The presence of water in the seed is also important as this is a medium
which readily absorbs energy. It seems probable, therefore, that the predominant mechanism causing the death of seeds and plants when high
power microwaves are used for weed control in a field situation is thermal.
IV.E (iii) Practical weed control by microwaves
Davis (1974) has listed more than seventeen species of weeds as well
as nematodes, fungal diseases and soil-borne insects, successfully controlled by microwave machines, with control lasting up to 12 months.
His experience and that of others clearly demonstrates that, in practice,
weeds can be killed by microwaves.
Values of 77, 800 and 1600 J cm-2 have all been quoted as the minimum

208
energy necessary for weed control (Hightower et al., 1974; Olsen, 1975;
Wayland et al., 1975). Variations in soil type, conditions and the desired
depth of treatment all influence the amount of energy needed. Energy
densities between 200 and 400 J cm-2 are commonly used. Values of 560
J cm-2 produced inadequate control of Johnson grass (Sorghum halepense), 2240 J cm-2 being required for good effect (Wayland et al., 1975).
The amount of energy required determines the speed of treatment as
well as the cost. If 200 J cm-2 is taken as a reasonable figure for energy
density and a 60 kW source is used, it can be calculated that the treatment
time is 92.6 hours per hectare. If only strips were treated this figure might
be reduced by a factor of three or four but it is still a long time. Hightower
et al. (1974) claim that energy densities of 1600 J cm-2 (i.e. 741 h ha-')
are required, and for control of Johnson grass (Sorghum halepense) (Wayland et al., 1975) an entire hectare would take 1037 hours. Lal and Reed
(1980) calculated that a 30 kW generator (2450 MHz) used to kill wild
oats (Avenafatua) in soil would cover only 1.8 x 10-3 ha h-I at an
operating speed of 0.1967 km h-1. The times taken are independent of
applicator area and can be reduced only by increasing generator power
or by reducing the treatment (spraying one hectare with herbicide should
not exceed one hour).
The Phytox Corporation was founded in 1971 to develop microwave
weed controlers (Phytox, 1971) and they produced several machines. The
biggest was the Zapper III, a 33,000 kg device. It used a 155 kW dieselalternator set to provide power for two 30 kW, 2450 MHz microwave
generators and the hydraulic drive system. This machine was used for
field trials and demonstration purposes. The Zapper IV was to be the
production model, having the same microwave generator power but some
10,000 kg lighter. It was to be a four-row field machine. Estimates of
working capability were from 0.4-2 ha daily per machine and it was
planned that 14,000-18,000 ha would eventually be treated by microwave
machines. Estimates for cost ranged from $312 ha-1 (1973) for treating
ground prior to planting peach trees (a 50 cm band to a depth of 30 cm
with two side bands of 110 cm to 5-7.5 cm depth) to $38 ha-' for general
control when treating strips to 5-7.5 cm depth (Scientific American, 1973).
The Zapper IV was never, in fact, produced and the entire project has
lain dormant. Recently plans have been suggested for another company
to revive it but no definite statements as to future operations have been
made (1980, Oceanography Int., pers. comm.).
The Phytox Zapper III beamed 60 kW of radiation towards the ground
and although the radiator was close to the soil surface the possibility of
considerable leakage existed. No figures were given as to field strengths
surrounding the machine when operating. Monitors which protected the
drivers would have turned off the main generators if radiation exceeding

ELECTROSTATIC
209
10 mW cm-2 reached them but were never activated. Bulk scattering

within the soil can occur which results in some radiation re-emerging
some distance from the machine, especially in view of the high incident
power. Passers-byor agriculturalworkersnearby can be exposed to the
microwaves.It is unlikelythat, with presentHealth and SafetyLegislation
within Britain and Europe, these machines would be allowed to operate
there without elaboratesafety shielding.
It seems, therefore,that, in view of the high costs associated with the
constructionof the machines and their operation,the problems of maintaininga uniformtreatment,theirvery slow rateof operationand possible
safetyhazards,the method is not viable at presentfor practicalagricultural
work. It is very successfulin controllingweeds, however, and has advantages over flaming,hoeing and sprayingin that it can control seeds buried
to severalcentimetersas well as surfaceseeds and plants.The only feasible
role for microwavesin agricultureat present seems to be specializeduses
where areas of ground may have to be cleared without treatment by
chemicals, e.g. research farms. In situations very sensitive to chemical
spillage or pollution (e.g. land adjacent to waterways),it does offer an
alternativemethod of weed control, although very slow and expensive.
V. The Effect of Electric Currents Applied
Directly to Plants and Soils
V.A SOILS, PLANTS AND APPLIED CURRENTS
Rawson and Le Baron (1906) investigatedthe effect of electricalstimulation on the growth of plants-by means of a zinc plate at one end and
a copper plate at the other end-in experimental areas in glasshouses
containing loam mixed with manure. The two plates (30-130 m apart)
were joined by a copper wire and the whole formed an electrochemical
cell with a potential of about 0.5 V and a currentof 0.4-15 mA. Earlier
maturity(one week)and largesize of lettuceswerereportedin comparison
to neighboring, untreated plants. Similar experiments with cucumbers
needed the electrode plate size reducedto avoid rankgrowth of the crop
foliage. Singh (1932) passed weak (unspecified)currentsthrough soil in
which seedlings of gram (Cicer arietinum) were grown. The root growth
in the electrified pot was smaller than in the control pot (no electric
current)but shoots were more vigorous. The electrifiedroots produced
tertiaryrootlets while the others did not.
Lutkova (1965) evaluated the effect of electric currentsof 6-100 mA
(2 x 10-10 to 8 x 10-6 A cm-2) through soil in both field and culture
experiments. Low currentsthroughoutthe growth period increased the
yield of potatoes by 15-30% with starch content of the tubers up by
1-2%. Tomato yield was enhanced by 20-35% and sugarbeet crops in-

210
Table VIII
Change in bacterial densities in various media as a result of electrical stimulations
(from Stone, 1909)
Current
strength
(mA)
Medium
Sewage contaminated
pond water
0.1
0.1-0.3
Sterilized water inoculated

with Bacillus megaterium
Sterilized water inoculated
with Pseudomonas
0.1-0.3
Treatment
time
Bacterial numbers ml-'

Control
Electrified
1 day
2 days
3460
3440
1 day
16 days
11,000
243,000
32,000
7,650,000
1 day
9 days
6000
50,000
15,000
5,210,000
143,000
1,470,000
143,000
86,590,000
44,000
109,000
radicicola
Milk
Soil (sandy loam)
0.3
0
27 hours
0.1-0.6
1 day
17 days
33,470,000
28,780,000
37,930,000
32,860,000
30 days
19,290,000
35,000,000
creased. Sugar contents of the plants increased by 0.3-1% and 0.3-0.4%

respectively. Electrification was reported to stimulate photosynthesis, inhibit oxidation processes during maturation and lead to a shorter growth
period and greater productivity.
Stone (1909) passed small electric currents (0.1-0.6 mA) through various media containing bacteria. The electricity was generated by copper
and zinc strips connected to form a galvanic cell with the media as electrolytes. Bacterial growth was generally stimulated except in the case of
bacteria in the soil where there was some decrease in the population but
much less than in the control. Table VIII shows some of the results.
Other experiments in which jars containing milk were charged electrostatically showed that small quantities of charge increased bacterial
activity while large ones decreased it. In addition positive charge was
shown to have a greater effect than negative charge.
Lazarenko et al. (1968) report work by Kravtsov who stimulated nitrogen-fixing bacteria in soil with currents of 0.1 A, whereas values of
10-40 A had a bacteriostatic effect.
Stern (1923) passed electric currents through Berberis nitens, B. vulgaris
var. atropurpurea,Biophytumsensitivum,Mimosa spegazini and M. pudica. Reactions varied depending upon the polarity of the stimulus and
the strength of the current but stimulation was usually stronger at the

211
negativeelectrode.The electrical(ohmic) resistanceof a plant determined

the value of the currentthroughit and its responseto a stimulatingvoltage.
The resistancevariedin differentregionsof a plant,e.g. in Mimosa leaflets
of the same leaf differedin resistanceand older leaves had higher resistances than the correspondingparts of youngerones.
Cholodny and Sankewitsch(1937) caused currentsof 10-7 to 10-6 A
to flow from the base to the apex of isolated oat coleoptiles (Avenasativa
var. SiegeshaferSvalof) and obtained an initial accelerationof growth
which subsequentlyslowed down and eventuallyreturnedto normalwhen
the currentwas switched off. Reversal of the currentusually caused retardationof growth. It was suggestedthat the currentsaffectedthe translocation of growth hormones throughoutthe plants. The durationof the
growthincreasedependedupon temperature-it lastedlongerat 16?Cthan
at 20?C. An increase in current during an experiment caused a further
shortincreasein growthrate. Similarexperimentswith rye (Secale cereale
var. Pektus)gave similar results.
Tomato plants (Lycopersicumesculentumvar. Scotia) were subjected
to electriccurrentsof 3 to 30 AAby Blacket al. (1971) for varyingperiods.
This resulted in linear growth increases of 5% to 30% and increases in
uptakeand leaf contents of potassium (15%),calcium (15%),phosphorus
(14%)and total nitrogenif the plant was negative to the soil. Makingthe
plant positive reduced the growth. When the electrode in the plant was
connected to the positive pole of the voltage supply, tissue damage occurredaround it, but not when the electrode was made negative.
Dixon and Bennett-Clark(1927) passed pulses of 50 Hz a.c. of varying
amplitudes and durations through 1 cm2 samples of ivy leaves (Hedera
helix) and measuredchangesin the electricalresistance.For smallerstimuli (100 V rms) the resistancerose by 5% maximum within 20 minutes
and then fell to normal. For heavier impulses (120 V rms, 0.1 s) the
resistancefell by 5%(10 minutes)and then slowly rose againto its original
value aftera further60 minutes. The temperatureand season of the year
affectedthe response.
Pea plants (Pisum sativum)35-45 cm high were used by Diprose et al.
(1978b). Stainless-steelwire was pushed throughthe stems 25 cm above
soil level and various currentspassed through.Below 100 ,A the currents
remained stable or dropped slightly. Above this value tissue damage
occurredand the currentsrose to new levels. Increases in voltage both
increased the currents and caused further rises until -5-10 mA. The
increases then became self-sustainingand currentsrose rapidly without
any further voltage increase to 25-50 mA whereupon the plants burnt
into two parts. During the experimentsdropletsof moisturewere seen on
the outside of the stem. Similarobservationsto those of Blacket al. (197 1)
were made concerningthe effect of the electrodeon the plant. A positive

212
THE BOTANICALREVIEW
polarity caused tissue damage and broke circuit continuity whereasnegative polarity caused foaming aroundthe entranceand exit points of the
electrode in the stem and maintained the currentflow.
V.B WEED CONTROL BY HIGH VOLTAGES
Slesarev(1973) and Slesarevet al. (1972) reportthe use of electricspark

dischargesboth for clearingweeds from fallow soil, and simultaneously
increasingthe availabilityof soil nutrients.Dischargesof 30 to 50 kV and
10-6 second duration were used on a range of young weeds [including
lambsquarters(Chenopodiumalbum),various types of Amaranthus,wild
hemp (Cannabis ruderalis),snakeweed (Polygonum bistorta),bindweed
(Convolvulusarvensis), tartar buckwheat (Fagopyrum tataricum), wild
oats (Avenafatua), and bristlegrass(Setaria)] and all were dead after 46 days. Increaseof either or both the voltage pulse strengthor duration
acceleratedthe death of the plants, as did treatinglargersections of stem.
The more plants treatedper dischargethe less the effect.The application
of between 5 and 25 kV to plants of "WhiteMary"(Chenopodiumalbum)
3-4 cm tall resultedin the cessation of transpiration,photosynthesisand
respirationby the third day. The dischargewas reportedto increase the
content of nitrate nitrogen and also available potassium in the soil, and
therewereno detrimentaleffectson soil structure(in termsof aggregation)
or microbiologicalfeatures.
Bayev and Savchuk (1974) investigated the causes of plant death due
to dischargesand eliminated the temperatureof the spark channel, the
pressureon the shockwavefront, optical radiationfrom the dischargeand
magnetic fields, concludingthat damage to the plant cells was caused by
highcurrentdensitiespassingthroughthe tissue. Svitalka(1976), however,
suggestedthat the temperatureof the sparkchannelcan causelocal damage
to the tissue and the shockwave is capable of destroyingcell walls and
coagulatingcytoplasm. He notes that, before the flashover, the conductivity currentsare responsiblefor cellulardisruption.Klimov et al. (1970)
showed that there was a thresholdvalue below which the conductivity of
plant tissue was constant and after a discharge it rose rapidly as cell
membranes lost their semi-permeabilityand vacuolar sap escaped into
the intercellularspaces.
Bayev and Savchuk (1976) studied the design of dischargeelectrodes.
They devised an electrode system which had non-linear electric fields
when a cylindricalstalk was presentresultingin a dischargeat 25-50 kV.
Withouta plant stalka linearfield distributionoccurredso that discharges
did not start until higherpotentials were used (e.g. 80 kV). The electrode
consisted of a pair of round plates 3 mm or less thick and 40-100 mm
in diameterwith a 'V' shaped notch in each plate. They were placed with

213
their edges facing, separated by 50-100 mm and with the notches opposite
each other.
The Lasco Corporation of America (Howe, 1977) has completed extensive field trials on post-emergence weed control by electric currents
and now produces equipment. An electric generator is carried at the rear
of a tractor with a transformer to provide 12-20 kV and control and safety
switching. The high voltage is supplied to an electrode, held above the
crop or to one side of it, which contacts the weeds. Destruction is by the
large electric currents forced through the plants by the high voltage, causing rapid heating and bursting of the cells. By use of systems with powers
ranging from 15-200 kW and a variety of electrode shapes and sizes,
successful results are claimed on broad-leaved weeds, grasses, shrubs and
small trees which are comparable to chemical methods in cost and effectiveness and have fewer adverse environmental effects (Bramblett, 1977).
Because of its manner of application, the electrical method is not selective
between weeds and crops unless there is a significant difference in conductivities with the desired plants having the highest resistance.
Dykes (1977) reports on clearing three weed species-cocklebur (Xanthium sp.), giant ragweed (Ambrosia trifida) and sesbania (Sesbania exaltata)-up to 2.5 m high-from around second year sycamore seedlings
0.6-2 m high. Current limited to 5 A was used at a speed of 4-5 km h'-i
with the electrode 1.3 m above ground level. An estimated 90-95% of
the broad-leaved weeds were dead five days later, with only slight damage
to the sycamore. After 60 days the tree seedlings showed no evidence of
being treated. Preliminary work (Chandler, 1978; Dykes, 1978) was also
reported on treating young cotton plants (four weeks old) when surrounded
by weeds. Initial results on crop and weed species indicate the need for
careful timing of the operation and adjustment of the voltage, but the
Lasco Corporation is continuing the research (1981, D. Johnson, pers.
comm.).
Dexter and Feight (1979) state that the method is limited to the control
of annual weeds only since the underground root systems of perennial
weeds are able to withstand the treatment and regenerate growth.
Wilson and Anderson (1981) evaluated the effectiveness of the Lasco
electric discharge system (EDS) on three weed species-kochia (Kochia
scoparia), redroot pigweed (Amaranthus retroflexus) and common lambsquarters (Chenopodium album)-growing in crops of sugar beet (Beta
vulgaris). The control of all weeds was 32%, 39% and 47% during 1977,
1978 and 1979 respectively, with the increase attributed to improving
equipment. No lasting damage was done to the crop, although some
individual leaves that had been inadvertently touched by the electrode
died but the beet itself was not harmed. In 1979, the sugar beet yields of
the EDS weeded plots compared to the handweeded ones were 76% (ko-

214
Table IX
A comparison of costs of three different types of weed control for three different
areas (after Kaufmanand Schaffner,1982). The electric weeder covering 60 ha

has been taken as a reference for cost as 1.000
Treatedarea (ha)
Type of weeder
Relative cost (ha-')
100
Electric
Roller
Re-circulator
1.000
0.560
0.374
500
Electric
0.280
1100
Roller
0.393
Re-circulator
0.243
Electric
Roller
Re-circulator
0.224
0.234
0.374
chia infested), 93% (common lambsquarters infested) and 89% (redroot

pigweed infested). If all the weeds are taken together, in 1979, three EDS
treatments gave a sugar beet yield of 99% as compared to the handweeded
control, two treatments 87%, one treatment 82% and no treatment gave
only 75%.
Kaufman and Schaffner (1982) compared the energy requirements and
costs of the EDS treatment with a herbicide roller and a re-circulating
sprayer both using a glyphosate preparation. Investment and running costs
were included although no account was taken of the comparative effectiveness on weed control or crop yield (Beta vulgaris). Table IX shows
that the EDS treatment was most expensive on small areas but it became
cheaper than the herbicide roller on areas above 210 ha. It was the cheapest
method of all on areas above 920 ha.
Vigoureux (1981, 1982) used a Lasco EDS machine to treat weed beet
in sugar beet (Beta vulgaris). The equipment was mounted on a 135 h.p.
tractor and produced a voltage of 8000-15,000 V. Average treatment rate
was 2 ha h-', with an average fuel consumption of 5.78 1 ha-1. He concluded that good treatment can be achieved in weed densities of 1005000 stems ha- ' (i.e. 98.0-99.9% control). Above this too much electrical
power is required and in an 18,000 stem ha- ' infestation only 24% of the
weeds were destroyed. Diprose et al. (1978b) treated electrically three
plant types: Chrysanthemum segetum, Sinapis arvensis, and Beta vulgaris
(bolting). Higer voltages resulted in higher currents and shorter destruction
times, e.g. Sinapis arvensis (1 m in height) took 2000 V rms 0.29 A and
147.8 seconds to burn in two, but at 4000 V rms the maximum current

215
was 1.34 A and treatment time only 13.2 seconds. The larger the plant
the higher the power needed to kill it. It was also found that to kill plants
in the field, two or three times as much power as for their glasshouse bred
contemporaries are needed, but not the ten times reported by Chandler
(1978).
Diprose and Benson (1980) and Diprose et al. (1 980a, 1980b) describe
trials with an electrothermal weed control device to control bolting and
weed beet in crops of sugar beet (Beta vulgaris). A 52 kW tractor drove
a 24 kW 240 V rms generator which fed a transformer giving 8.4 kV rms
output. The electrode consisted of three metal rods 3 m wide, to span six
crop rows, placed one behind another to form a grid 1 m long, and it was
suspended hydraulically 8-15 cm above the crop. The whole machine
could move at 1.6 km h-I giving a plant/electrode contact time of 2 or
3 seconds. An interval of four weeks was left between the two treatments
of the trial plots and results were noted five weeks later. Of the 186 annual
beet counted in the trial plot, 51 had grown after the treatment but did
not produce viable seed. Of the remainder, in 101 the stems and leaves
were killed and no seed was produced. The 25% of treated plants that
had survived were those which grew along the top of or beneath the crop
rather than above it and so were missed by the electrode. A new machine
was constructed in 1981 using a 100 h.p. tractor providing an electrical
output of 54 kW at 9, 13 or 17 kV. It could travel at speeds in excess of
5 km h-1, spanned 12 rows of sugar beet crop, achieved a working rate
of 2.4 ha h-I and used between 3 1 ha-l and 6 1 ha-1 of diesel fuel in
medium infestations of weed beet (Diprose and Benson, 1982).
V.C
CONCLUSIONS
V.C(i) Summary
Small electric currents of 0.1 to 100 mA have been passed through soils
(Lazarenko et al., 1968; Lutkova, 1965; Rawson and Le Baron, 1906)
with reported beneficial effects to plants and bacteria. Heavy current of 40
A through soil was toxic to bacteria (Lazarenko et al., 1968). Stone (1909)
passed small electric currents (0.1-0.6 mA) through media containing
bacteria and stimulated their growth.
Small currents (10--_10-2
A) passed through plants and leaves (Black
et al., 1971; Cholodny and Sankewitsch, 1937; Dixon and Bennett-Clark,
1927; Stem, 1923) produce a variety of responses and can affect the
electrical resistivity of plant tissue, growth rates, and movement of metabolites. Heavier currents [i.e. 50 mA and above (Diprose et al., 1978c,
1980b; Howe, 1977)] cause the destruction of plant tissue by the rapid
heating of cellular water, leading to cell membrane rupture.
Weed killing machines with various types of electrodes have been con-

216
structed. Some use very high voltage discharges (e.g. 50 kV) (Bayev and
Savchuk, 1974, 1976; Slesarev, 1973; Slesarev et al., 1972; Svitalka, 1976)
and some use high voltage electrodes (8-20 kV) which have to touch the
plant being treated (Diprose and Benson, 1980; Dykes, 1977; Kaufman
and Schaffher, 1982; Wilson and Anderson, 1981). Machines of the first
type are complicated and can deal with small plants only (a few centimeters
tall) but the direct contact machines are more powerful and can handle
large weeds and heavy infestations.
V.C (ii) Practical weed control by electricdischargesand currents

The best method of weed control using high voltages is the direct contact
rather than the pulsed discharge method. The voltages are lower so there
are fewer operating and insulation problems- 12-20 kV as opposed to
30-50 kV-and the generating and control circuitry is simpler and cheaper. The electrodes are simpler-rods spanning the crop rows-and can
touch a variety of weed types as well as handle a range of densities at
once. The Russian discharge system can treat only one or two small plants
at a time and that is provided the round electrodes can get to both sides
of the target plant. The pulsed discharge treats plants a few centimeters
tall whereas the direct contact can deal with those 2.5 m high or more.
Experience in Europe and America with the direct contact method
indicates that high voltages (12-20 kV) are necessary for fast speeds through
the crop, i.e. 5 km h- 1, and high powers are required (i.e. 50 kW or more),
to cope with large weeds or large infestations of small ones.
Both pulsed and direct contact types are only post-emergence systems
and, except in a few circumstances, the weeds must be separated in some
way from the desired plants. It is not therefore a general replacement for
herbicides but useful where a late weed outbreak occurs or where herbicides cannot be used, e.g. bolting sugar beet. A number of weeds can be
missed. For example, Wilson and Anderson (1981) report only 47% of
weeds treated in sugar beet in 1979 and Diprose et al. (1 980a) report 25%
of annual beet in sugar beet missed. This is the same order, however, as
mechanical cutting, herbicide rollers or re-circulating sprayers. Kaufman
and Schaffner's cost analysis indicates that electric weed control becomes
cheaper than other methods when the areas to be treated are large.
The machines now marketed by the Lasco Corporation have several
features to ensure the safety of the operator. There is an off switch in the
operator's seat so that the high voltage is switched off should he or she
move or fall from the normal position. Other switches are activated and
turn off the electrical system if parts of the equipment overheat, earth
continuity is broken, the tractor is moving at less than 1.6 km h-I or the

ELECTROSTATIC
217
generatingequipmentis lifted on the hydraulicswithout it being switched

off.
Europeanmachines have similar devices but regulationsalso require

extensive shieldingof the active electrodeto minimize dangerto bystanders. In GreatBritainit is thoughtthat the machinewill only be authorized
for use as an agriculturalcontractor'stool and that shorttrainingprograms
for operatorswill be required.
Acknowledgments
The authors thank the Council of the Perry Foundation for financial
assistance duringthe period of the preparationof the manuscript.
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Effects of Electrostatic Fields, Microwaves and Electric Currents on Plants

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The Effect of Externally Applied Electrostatic Fields, Microwave Radiation and Electric

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The Effect of Externally Applied Electrostatic Fields,

The Botanical Review 50: 171-223, April-June, 1984

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THE BOTANICAL REVIEW

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ELECTROSTATIC FIELDS, MICROWAVE RADIATION

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THE BOTANICAL REVIEW

Mechanical methods of removing weeds, e.g. hoeing and tilling, have

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weed seeds of the type that can withstandhigh temperatures,e.g. as great

An electrostatic field is an electric field that is static in time, i.e. its

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Microwaveradiationis the name given to electromagneticradiationof

where f is the frequencyin Hz; E the field strengthof radiation at point

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II.C ELECTRICAL DISCHARGES AND DIRECT ELECTRIC SHOCKS

Electricdischargesin the context of this review are taken to mean the

A number of studies of the effects of electric fields on the growth of a

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to the demonstrationsthat growth can be stopped and leaves and whole

PLANT GROWTH IN THE PRESENCE OF ELECTRIC FIELDS

Earlyresearchsuggestedthat the effects of subjectingplants to electric

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(percentage of control samples)

field strengths of 30, 50 and 75 kV m-1 to seedlings of orchard grass (in

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ELECTROSTATIC FIELDS, MICROWAVE RADIATION

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r damagewas almost instantaneous(afteronly a few seconds

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ELECTROSTATIC FIELDS, MICROWAVE RADIATION

Bankoskeet al. (1976) investigatedthe effectsof 60 Hz electromagnetic

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THE BOTANICAL REVIEW

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(Kotakaet al., 1965) as comparedto controls. Increasesin air ion density

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III.D (i) Summary

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ELECTROSTATIC FIELDS, MICROWAVE RADIATION

1968) can be increased by air ions. Barley seedlings grown in ion-depleted

III.D (ii) Practical weed controlby high electricfields

Microwaves and Weed Control

IV.A USES OF MICROWAVES IN AGRICULTURE

Microwave radiation has been suggested as a possible solution to many

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which was sufficientto maintainthe leaf temperatureat 25?C.The authors

1.5 cm of snow. A 2.4 kW generatorwas used and the irradiatingantenna

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when wood temperaturesof 50?Cwere obtained. They predicted that a

temperaturewas found to rise from 200Cto nearly80?Cwithin 2 minutes

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THE BOTANICAL REVIEW

non-aqueous volatile substances. Other authors (Hankin and Sawhney,

ON PLANTS AND SEEDS

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ELECTROSTATIC FIELDS, MICROWAVE RADIATION

Phaseolus linensis vars. Hendersonbush and Fordhook242, Stizolobium