Microwave Radiation and Plants

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    The Effect of Externally Applied Electrostatic Fields, Microwave Radiation and ElectricCurrents on Plants and Other Organisms, with Special Reference to Weed ControlAuthor(s): M. F. Diprose, F. A. Benson and A. J. WillisReviewed work(s):Source: Botanical Review, Vol. 50, No. 2 (Apr. - Jun., 1984), pp. 171-223Published by: Springeron behalf of New York Botanical Garden PressStable URL: http://www.jstor.org/stable/4354034.

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    T H

    BOT NIC L R V I W

    VOL.

    50

    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,University f

    Sheffield

    Mappin

    Street, Sheffield,SI 3JD United

    Kingdom

    AND

    A. J. WILLIS

    Department

    of Botany, University f Sheffield,

    WesternBank

    Sheffield,

    S1O

    2TN

    UnitedKingdom

    Abstract

    .----------------------------------------172

    Sommaire.------------------------------------------------173

    I. Introduction

    ..14...

    ..............

    ....

    174

    II. Definition

    of Forms

    of

    Electrical Energy

    ..

    . 175

    A.

    Electrostatic

    Fields

    .---------------

    .----.--..--..-----175

    B.

    Microwave Radiation

    .

    176

    C.

    Electrical

    Discharges and Direct Electric

    Shocks

    .

    177

    III.

    Electrostatic

    Fields

    and

    their Lethal Effects

    on

    Plants

    ..-------------------------------------------

    77

    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

    ..18------------------------------------------------------------------------------------------------------86

    (i) Summary

    186

    (ii)

    Practical Weed

    Control

    by

    High

    Electric

    Fields

    ........................................ ......

    187

    IV. 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

    and

    handling

    fee. Valid until 31 December

    1984):

    U.S.

    Orders:

    $10.75.

    Non-U.S.

    Orders:

    $11.50.

    (Payment,

    U.S.

    currency

    only

    and either

    drawn

    on

    a U.S. bank or

    made

    out

    by

    intemational

    money order,

    should

    accompany

    order. Thank

    you.)

    The

    Botanical

    Review

    50: 171-223,

    April-June, 1984

    171

    ?

    1984

    The

    New

    York Botanical

    Garden

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    172

    THE BOTANICAL

    REVIEW

    C. Field Experiments

    with

    Microwaves for W eed Control

    ................................................96

    D. The Effect of

    Microwave

    Radiation

    upon

    Soil

    Microorganisms

    and

    N em ato des ..----

    --- --

    ---

    ---

    --

    ---

    --- -- ---

    --- -----

    --- ---

    --

    --- ---------- ---

    202

    E. Conclusions ..204------- . ................. 204

    (i)

    Summary.

    -------------------------------------------

    -----

    204

    (ii)

    Thermal or

    Non-thermal Mechanisms

    of Death 206

    (iii)

    Practical

    Weed Control

    by

    Microwaves .................

    207

    V. The Effect of

    Electric

    Currents

    Applied Directly

    to

    Plants

    and Soils

    .

    209

    A.

    Soils,

    Plants and

    Applied

    Currents

    .

    .....

    209

    B. Weed Control

    by

    High Voltages

    ...........

    212

    C. Conclusions

    .............................................................

    215

    (i) Summary

    ----------------------------------------------------215

    (ii) Practical Weed Control by Electric Discharges

    and Currents

    ......

    216

    VI. Acknowledgments ................ 217

    VII. Literature Cited.

    -----------------..----------------------------------------------------------------------------

    17

    Abstract

    A

    wide-ranging

    review is

    presented

    of

    the effects

    of

    various forms of

    externally applied

    electrical

    energy upon plants

    and

    other

    organisms.

    Al-

    though 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.

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    ELECTROSTATIC

    FIELDS,

    MICROWAVE

    RADIATION

    173

    Electric currents

    have been

    caused to

    flow

    throughplants

    by

    the

    ap-

    plication of

    electrodesto

    the

    leaves. The effects

    range

    from

    nil,

    when

    50-

    100 V and 1 or 2 ,uAare used, to very strikingwhen voltages from 5 to

    15 kV

    areapplied

    causing

    currents

    of several

    amperes

    o flow and

    resulting

    in therapid

    destructionof

    the

    target.Smallelectric

    currents

    passed

    through

    soil

    containingplants

    are

    reported

    o increasetheir

    growth.

    The

    effects

    of

    small current on the

    growth

    of

    individual leaves are reviewed.

    The

    use

    of

    high

    voltage

    tractor-borne

    quipment

    for

    weed control is

    also consid-

    ered.

    Sommaire

    Une revuea

    larges

    themes

    presente

    les effets

    des diverses

    formes

    d'ap-

    plication externe

    d'energieelectriquesur les

    plantes

    et

    autres

    organismes.

    Bien

    que

    desrecherches

    omportant

    a

    la fois de

    petites

    et

    grandesquantites

    d'energie

    dirigees

    sur les cibles en

    question

    soient

    prises

    en

    consideration,

    un

    des

    aspects

    particuliers

    de

    cette

    revue est

    la

    possibilite

    d'application

    de

    chaquetype de

    stimulation

    electriqueau

    controle

    des

    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'.

    Quelques effets

    benefiques

    ont ete

    enregistres,

    par

    exemple, l'accroissement

    de

    la

    production

    des

    recoltes

    de

    cereales

    et

    de

    legumes; mais les

    resultatsetaient

    irreguliers

    et les

    conditions

    elec-

    triques

    conduisanta des

    profits

    bien

    determines

    a grandeechellene

    pou-

    vaient

    pas

    etre

    predites avec

    confiance

    des

    premieres etudes.

    On s'est

    apercu

    que

    de

    grands

    champs

    electriquespouvaient deteriorer

    es

    plantes

    si des courants

    superieurs

    a 10-6 A

    etaient

    amenes

    a

    circuler

    dans

    les

    feuilles, entrainant

    des

    dechargesde la

    couronne

    a

    partirdes

    pointes. La

    nature des degats ainsi que les effets sur les procedes metaboliquessont

    ici

    etudies, de

    meme

    que les

    resultats

    des

    experiences sur

    la

    croissance

    des

    plantes

    pour lesquelles la

    densite et

    la

    chargedes ions

    dans

    I'airont

    ete

    changes.

    Les

    effets du

    rayonnement

    par

    micro-ondes

    (pour

    la plupart

    de

    2450

    MHz)

    sur

    les

    graines,

    les

    plantes

    et

    autres

    organismesdu sol

    sont

    ici

    exposes. Ces

    effets

    dependent

    de la

    densite electriquedu

    rayonnementet

    des

    proprietes

    electriquesdes

    objectifs.

    Des

    facteurstels

    que la

    taille des

    graines et des plantes,leur forme et leurteneuren humidite sont impor-

    tants, comme le

    sont les

    proprietesd'un sol

    irradie

    (notamment

    sa

    teneur

    en

    eau).

    Bien

    que les

    micro-ondes

    puissent

    etre efficaces

    pour

    tuer des

    plantes et aussi

    des

    graines

    enterreesa plusieurs

    centimetres de

    profon-

    deur,un

    important

    appareillage

    electrique est

    necessaire

    et

    les

    temps de

    traitement

    sont

    longs; par

    exemple une

    machine de 60 kW

    peut

    prendre

    jusqu'a

    92,6

    h

    ha-'.

    D'autres

    experiences

    demontrerent

    que le

    rayonne-

    ment

    par micro-ondes

    pouvait

    tuer les

    nematodes

    dans la terre

    et qu'il

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  • 7/26/2019 Microwave Radiation and Plants

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    174

    THE

    BOTANICAL

    REVIEW

    etait tres

    efficace

    pour

    detruire

    es

    champignons

    et les

    bacteries

    du

    sous-

    sol.

    Le

    potentiel

    des

    diverses

    utilisations

    possibles

    de la

    radiation

    par

    micro-ondes en agriculture st egalementdecrit.

    Des courants

    electriques

    ont

    ete

    amenes a

    circuler

    a

    traversdes

    plantes

    par

    l'application

    d'electrodes

    sur

    les

    feuilles. Les

    effets,

    nuls

    quand

    50

    a

    100

    V

    et

    1

    a

    2

    ,uA

    sont

    utilises,

    sont

    par

    contre

    frappants

    quand

    des

    voltages de 5 a

    15

    kV

    sont

    appliques,entrainant

    a

    circulationde

    courants

    de

    plusieurs

    amp'eres

    et la

    destruction

    rapide

    des cibles. On

    remarque

    cependant

    que,

    de

    petits

    courants

    electriques

    envoyes

    dans un

    sol conte-

    nant des

    plantes, accelerent

    eur

    croissance;

    es

    effets d'un

    faible

    courant

    sur acroissancedesfeuilles ndividuellessontici 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

    practiced

    for

    centuries

    (Crofts,

    1975).

    The

    use of

    chemical

    methods

    for weed

    control

    has

    been

    expanding

    at

    a fast

    rate

    since the

    mid

    1940's

    (House,

    1967).

    In

    recent

    years electrical

    methods have

    been

    investigated

    in

    this

    connection,

    some of

    which

    promise

    to be of

    practical

    value. This

    review

    discusses

    the

    control of

    weeds,

    and

    also of

    plant

    growth

    more

    generally,

    by

    means of

    such

    methods.

    Weeds can

    be

    subjected o

    electrical

    energyby use of

    electrostatic ields,

    microwaves,

    electric

    discharges

    or

    direct

    electric

    shocks

    using either

    al-

    ternating

    current

    (a.c.)

    or

    direct

    current

    (d.c.). These

    techniques

    have

    several

    advantages

    over

    present

    methods of

    weed

    control

    and,

    although

    they cannot be regardedas a panaceafor all problems,electricalmeans

    appear to offer

    viable

    and useful

    additions to

    the

    stock

    of

    weed

    control

    methods.

    During

    and

    following

    the

    applicationof an

    electric

    shock

    or

    microwave

    or

    laser

    irradiation, he

    energy,

    rapidly

    absorbedby

    the

    "load,"

    is

    largely

    dissipated

    as

    heat

    in

    the

    plant.

    No

    residue is

    left to

    contaminate

    the soil,

    an

    important

    feature

    n

    view of

    rising

    concern

    about

    the

    contributionby

    herbicides to

    environmental

    pollution.

    The

    electrical

    energy

    can also

    be

    directedto where it is required,even underfairlyadverseweathercon-

    ditions.

    Electrodes

    carrying he

    currents,

    or

    the

    radiation

    being

    directed

    towards

    the

    ground

    are,

    unlike

    sprayed

    herbicides,

    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

    centimeters

    depending

    upon

    the

    applied

    power,

    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

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    ELECTROSTATICIELDS,MICROWAVE

    RADIATION

    175

    weed seeds of the

    type

    that

    can

    withstand

    high temperatures, .g.

    as

    great

    as

    1

    270C

    f

    slightly

    below the surfaceor if in cracks n the soil

    (Sampson

    and Parker,1930). Microwavepower would reach these seeds and still

    be as effective

    in

    killing

    them as if

    they

    were on the

    surface.

    While electrical

    methods

    of

    weed

    control are a

    relatively

    recent inno-

    vation, some

    forms

    of electrical and

    electromagneticenergy

    have

    long

    been

    known to be

    essential

    in

    agriculture-notably electromagneticra-

    diation

    in the visible

    wavelength region

    (Borthwick, 1965)

    necessary

    for

    photosynthesis.Other

    plant processes

    in which

    light

    is involved

    include

    photoperiodismandgermination.

    Furthermore,higherfrequency

    electro-

    magneticwaves such as y-rays may be importantin evolution, leading

    to the formation

    of

    new

    species, by

    causing

    mutationswhen absorbed

    e.g.

    by seeds

    (Nelson, 1965).

    The naturalelectricalstate

    of

    the

    atmospheremay

    be

    important

    o

    plant

    growth (Wheaton, 1970). The air contains both

    positive and

    negative

    ions,continuouslyprovidedby

    radioactivedecay of elements

    n

    the earth's

    surfaceand

    some

    cosmic radiation

    (Chalmers, 1967).

    Ion densities

    vary

    throughout

    he

    day

    but

    over

    open

    land are

    in

    the

    order of 1.5-4.0

    x

    103

    ions cm-3.

    Kotaka and Krueger 1968) have examined the effectsof

    vary-

    ing ion densities

    of air

    on

    the growth of

    barley, oats and lettuce. They

    report that increases

    n

    ion density lead to

    vigorous, accelerated

    growth.

    Alternativelyplants exposed to

    ion-depleted atmospheres ack vigor

    and

    have soft

    leaves (Kruegeret al., 1965).

    The effects of electrical

    energy are therefore

    mportant

    to

    plant life

    in

    many ways and

    electromagnetic adiation s vital to it. For weed

    control,

    however, larger

    than normal applicationsof electrostaticfields,

    electric

    discharges,and

    microwave radiationto plants

    are necessary.Thesetreat-

    ments arereviewed below, following a short explanationof the technical

    definitions of the

    various

    forms

    of electrical

    energy.

    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

    betweendifferent

    parts

    of the

    plant (Lund,

    1947). Such naturallyoccurring

    phenomena

    are

    not

    considered

    n

    this

    review which deals with the various

    waysin whichplants can be subjected o electricalenergy rom an

    external

    source-typically many

    orders of

    magnitudegreater han internally

    gen-

    erated

    electrical

    effects.

    II.A

    ELECTROSTATIC FIELDS

    An

    electrostatic field is an

    electric

    field that is static

    in

    time, i.e. its

    strength

    does not

    vary through

    time

    (Nussbaum, 1966). The field can be

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    176

    THE

    BOTANICALREVIEW

    formedby 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 de-

    fined as

    V

    E

    =

    d

    where E is the electric field strength

    n

    the air gap between the

    electrodes

    (conductors)

    measured

    in

    volts

    per meter,

    V

    the

    voltage applied

    to the

    conductors

    n

    volts,

    and

    d the distance between

    them

    in

    meters. For the

    parallel-plateconfiguration he field is uniformbetween the conductors,

    but

    differentexpressions

    exist

    to

    describe other more

    complex

    non-uni-

    form

    field patternsthatarise,

    for

    instance,

    with one

    planar

    electrode

    (e.g.

    earth)

    and an overhead

    wire,

    or when a

    plant

    is

    introducedbetween

    two

    parallelplates.

    II.B

    MICROWAVE

    RADIATION

    Microwave radiation s the name given to

    electromagnetic adiationof

    which the wavelengthsare

    much

    greater

    han

    those

    of

    light,

    i.e.

    they

    are

    measured

    n

    mm

    or cm

    (Glazier

    and

    Lamont, 1958).

    A

    wavelength

    com-

    monly

    used

    in

    weed control s 12.25

    cm which

    corresponds

    o a

    frequency

    of 2450

    MHz.

    Most of the

    papers

    referred

    o

    in

    this review that describe

    microwave

    experiments

    use this

    frequency.

    This is one of

    the

    frequencies

    allocated, by

    international

    agreement,

    to

    microwave

    power

    devices

    for

    domestic and

    industrial

    applications.

    A

    wide

    range

    of

    equipment

    s

    com-

    mercially

    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

    frequency

    n

    Hz;

    E

    the

    field strength

    of

    radiation at point

    of

    absorption

    n

    Vm-

    1; r"

    is the relative

    dielectric oss factor(Nelson and

    Wolff, 1964; White, 1970).

    Thus

    the power absorbeddepends upon several

    factors ncludingprop-

    erties of the load itself such as the relative dielectricloss factor,and the

    homogeneity of the medium. The electric field

    strengthat any point in

    the

    load

    depends upon

    the

    load

    shape and the

    dielectric

    properties

    of

    the

    medium,

    as well as the

    strength and frequency of the irradiating ields.

    The radiation

    penetratesall parts

    of

    the load

    simultaneouslywhich is

    in

    contrastto the normal

    heating process where heat

    is conducted from the

    outside

    of

    the load towards the

    center. The

    speed

    of action is

    one of the

    main

    benefits of microwave radiation.

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    ELECTROSTATICIELDS,MICROWAVE

    RADIATION

    177

    II.C ELECTRICAL

    DISCHARGES

    AND DIRECT ELECTRIC

    SHOCKS

    Electricdischarges n the context of this review are takento mean the

    discharge

    of

    a

    quantity

    of

    electricity

    into

    the

    plant

    from

    a

    high voltage

    electrical

    energy device,

    usually,

    but not

    necessarily,

    a

    capacitor

    (Duffin,

    1965). When the electric

    charge

    s

    pulsed,

    the

    generator

    has to

    chargeup

    a capacitor,and when

    that

    operation

    s

    completed

    the

    energy

    s

    transferred

    to the load. Repetition

    rates

    depend upon

    the

    generatorsize,

    the

    storage

    capacitance

    and the

    properties

    of the load under

    discharge.

    Russian re-

    searchers

    have

    used

    this method

    with

    up

    to 60

    kV

    pulses

    of 1

    As

    duration.

    The discharge

    electrodes need

    not

    necessarily

    touch the load

    if

    very high

    voltages are used, since

    the

    discharge

    can

    jump

    across

    an

    air

    gap.

    Alter-

    natively, a pulsed high voltage

    discharge

    can

    be

    applied

    from

    the

    sec-

    ondarywindingsof a high

    voltage

    transformer,

    without

    using

    a

    capacitor,

    by

    a

    suitable

    control of the

    input voltage

    to

    the

    primarywindings.

    Direct electric

    treatment is a continuous

    process whereby

    a

    generator

    is

    physically

    connected to the

    plant by

    two

    electrodes,

    or

    by

    one

    electrode,

    the

    ground

    being

    used as the other. The

    voltage

    can

    range

    from 500

    V

    to

    several

    kilovolts as

    required,

    and can

    be

    a.c.

    or d.c. The

    current,

    I

    (A),

    flowing throughthe plantis determinedby its electricalresistance,R (Q),

    and

    the

    applied

    voltage,

    V

    (volts),

    so

    that

    V

    R

    The

    units of alternatingvoltage and

    current

    are

    volts

    or

    amps r.m.s. The

    latterterm standsfor"rootmean

    square"and describes

    heeffectivevalue

    of

    a

    voltage

    or current

    waveform that varies

    continually

    with time

    (e.g.

    an alternatingcurrentwhich has the same effecton a load as a 1 A direct

    current

    has

    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 attributed

    to electrical

    action. Shlantaand Moore

    (1972) have also

    observed

    damage (browned

    eaf tips) in the

    "natural" rassof a mountain

    meadow

    in

    New

    Mexico

    after

    storms. Suchobservations

    have led to the

    investigationof the

    possibilities of a "lethal

    electrotropism"n plants and

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    178

    THE

    BOTANICALREVIEW

    to

    the

    demonstrations

    that

    growth

    can

    be

    stopped

    and leaves and

    whole

    plants killed

    by

    higher

    electric

    fields.

    III.B

    PLANT GROWTH IN THE

    PRESENCE OF

    ELECTRIC FIELDS

    Earlyresearch

    suggested

    hat the

    effects

    of

    subjectingplants

    to

    electric

    fields were beneficial.

    This

    topic

    has been the

    subject

    of research

    and

    debate

    for

    many years(Ellis

    and

    Turner,1978;Pohl,

    1977;

    Sidaway, 1975;

    Wheaton,

    1970),

    but the

    beneficial effect

    is

    not now

    generallyaccepted.

    Fromthe mid

    eighteenth

    o

    the

    early

    twentieth

    century,

    indings

    ndicated

    that increasedyields from both cereal and vegetablecropscould be ob-

    tained by

    applying electrostatic

    fields to

    plants

    while

    they

    were

    growing

    (Hendrick,

    1918; J0rgensen

    and

    Priestley,

    1914;

    J0rgensen

    and

    Stiles,

    1917;

    Newman, 191 1; Shibusawaand

    Shibata,

    1930).

    Between 1885

    and 1903

    Lemstrom(1904) undertook

    various

    experi-

    ments on the

    effects of

    electrical fields on

    plants

    in

    different

    places

    in

    Europe-from Finland

    and Sweden

    n

    the

    north,

    to

    England

    and

    Burgundy

    in

    the

    south.His

    results,

    while

    encouraging,

    were not

    very

    consistent

    (one

    factor

    being

    the

    unreliability

    of

    earlyhigh

    voltage

    equipment).

    His

    general

    conclusion was that electricfields, appliedto cropsby wiresstrungabove

    the

    growing

    areas,

    were

    beneficial,

    producinghealthy

    plants

    with

    increased

    yields.

    This applied

    to root

    crops,

    vegetables,cereals and

    strawberries.

    He

    proposed

    that the best

    times for

    applying

    the

    electrostatic

    field

    were

    for

    four

    hours

    in

    the

    early

    morning

    and

    another four

    during

    the

    late

    afternoon.

    The

    electricity

    could be

    applied

    all

    day

    duringcloudy weather

    and

    during

    nights

    of

    moist weather.

    Application

    duringdry conditions

    and

    in

    strongsunshine,

    however,

    could have

    adverse effects.

    Lemstrom'sstudies were extendedby severalworkers.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

    showedincreased

    yields;

    9 of

    these increases

    were of

    at

    least

    30%.

    In

    pot-culture

    experiments

    carriedout on

    maize,

    wheat and

    barley,

    the

    average

    yield

    increase,

    when

    these

    cereals

    were

    subjected to

    weak

    electric

    currents,

    was over

    11%

    (Blackman

    and Legg,

    1924). The

    Board

    of

    Agriculture

    and

    Fisheries of

    the

    U.K. even

    established a

    Com-

    mittee forElectroculturen 1918;however, its finalreport n 1937 (Board

    of

    Agriculture

    and

    Fisheries,

    1918-1937)

    stated

    that the results

    of 18

    years

    of

    research

    were not

    conclusive

    and it

    was not

    possible

    to predict

    confidentlythe

    benefits of

    electrical

    treatmentsto

    crops.

    The

    encouraging

    results of the

    European

    workers

    were not,

    however,

    shared

    by

    their

    American

    colleagues,who, in

    two

    fairly

    comprehensive

    sets

    of

    controlled

    trials, found no

    significant

    beneficial

    effects of electric

    fields

    on

    growth

    (Briggs

    et al.,

    1926;Collins et

    al.,

    1929). In view

    of these

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    ELECTROSTATIC

    IELDS,MICROWAVE

    RADIATION

    179

    findings

    and the

    inconsistency

    of other

    results,

    the

    use

    of

    electricaltreat-

    ments of plants lost

    favor.

    Apart

    from

    a few

    supporters

    Sidaway,

    1969;

    SidawayandAsprey,1968),the methodsdid not attractattentionalthough

    more

    recently

    interest

    has

    been

    shown

    in

    the

    effects of air

    ions,

    rather

    than electric

    fields,

    on

    plant growth

    (Bachman

    et

    al.,

    1971;

    Kotaka

    and

    Krueger,1968;

    Kotaka et

    al., 1965; Krueger

    et

    al., 1965,

    1978).

    III.C THE LETHAL

    EFFECTS

    OF

    ELECTRIC

    FIELDS ON PLANTS

    The term "lethal

    electrotropism"was

    suggestedby

    Murr

    (1964a)

    fol-

    lowing a

    series

    of

    experiments

    on

    the effects

    of

    electric fields on

    plants.

    Intheseexperiments Murr,1963a, 1963b)anelectric ieldwasestablished

    between two aluminium

    wire

    grids

    (0.24 m2).

    A

    lower

    electrode

    was

    situated below the

    soil

    in

    a plot

    in which

    seedlings

    of orchard

    grass

    (Dac-

    tylis

    glomerata)

    were

    planted.

    An

    upper

    electrode was

    suspended

    above

    the soil and

    adjusted

    n

    height

    to

    vary

    the

    electric field

    strength,

    but

    was

    never

    more than 10 cm above the

    tops

    of

    the

    plants.

    Temperature

    and

    light

    intensity

    werecontrolled

    (unspecified)

    and a 16-hour

    day length

    was

    used.

    The

    control

    plots

    had

    the same

    electrode

    arrangement

    s the

    "active"

    ones, but withoutthe voltagesapplied.The top electrodewas madepos-

    itive and

    the

    bottom

    one

    connected to the

    negative

    of

    the

    power

    supply

    simulating

    the

    earth's natural

    electric field

    (Chalmers,1967).

    Murr

    (1963a,

    1963b)

    observed

    that

    during

    continuous

    exposure

    to the

    electrostatic ields the leaf

    tips

    of

    the

    seedlingsbegan

    to

    brown,

    as if

    burnt,

    and

    he noted the

    similarity

    to

    mineral

    deficiency

    symptoms.Between

    the

    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

    downwards romthe tipat a fasterratethan thegrowthof theplant(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 ob-

    tained. Murr

    defined a

    damagefactor as the

    proportion

    of

    the

    dry

    weight

    of

    electrifiedto

    control

    samples

    (averageof three

    results)

    expressed as a

    percentage.

    For

    orchard

    grass,

    damage was found to rise

    to

    25%at a field

    strength

    of 50

    kV

    m-1

    and

    then

    rapidly increasedto

    50%

    at

    75

    kV

    m-l.

    Similarresults

    were

    obtained with

    seedlings

    of

    reed

    canary

    grass(Phalaris

    arundinacea) Murr,

    1963b).

    Transversesections

    of some

    leaves showed

    that the epidermalcells had been destroyed n the

    brownedtip and

    dam-

    aged

    in

    the dark

    green

    zones. There

    was

    completeabsenceof cell

    structure

    in

    the

    tip area

    and chloroplast

    derangement n the

    darkgreen

    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

    resulting

    concentrationunbalance

    might

    upsetnormal

    osmotic

    phenom-

    ena and

    cause

    rupture

    of the

    cells.

    To

    investigatethis

    further

    he

    applied

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    180 THEBOTANICAL

    REVIEW

    Table I

    The

    concentrationof elements

    n

    the

    leaftipsof

    orchard

    grass Dactylis

    glomerata)

    seedlings afterexposure to electrostatic ields(after Murr,1964b)

    Electrostatic

    strfielgth

    Phosphorus Nitrogen

    Iron

    Zinc

    Aluminum

    (kV

    m-')

    (percentage of control

    samples)

    30

    104

    95

    119

    177

    104

    50

    94

    97

    169

    258 233

    75

    97

    98 202

    317

    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

    (1964a,

    1964b)

    concluded that

    the

    lethal

    damage

    was not

    caused

    by

    the

    drift of the

    ionized

    salts, leading

    to the

    bursting

    of

    cells

    and

    de-

    hydration,

    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), experi-

    menting

    with

    oats,

    reported that,

    after one

    month,

    the electrified

    crop

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    ELECTROSTATIC

    FIELDS,

    MICROWAVE

    RADIATION

    181

    was taller

    and greener

    than oats

    not

    subjected

    to

    electric fields.

    Again

    in

    an

    experiment at

    Rothamsted

    Experimental

    Station

    in

    1917,

    Blackman

    (1924) foundthatbarleywas tallerandgreenerafter20 daysof application

    of the electric

    field

    than that

    in the

    neighboring

    control

    plot,

    although

    the

    visual

    differencebetween

    the

    crops

    subsequently

    became

    less

    marked.

    Priestley

    19 10)

    suggested without

    proof)

    that the

    deepergreen

    n

    wheat

    might result

    from

    a

    slight

    but continuous amount of nitrates

    being

    added

    to the

    soil

    by

    the overhead

    discharge,perhaps

    ormed

    in

    a similar

    manner

    as the combination of

    oxygen

    and

    nitrogen produced

    by

    thunderstorms

    is

    washed

    into

    the

    soil by rain. One soil test

    (Priestley, 1907)

    indicated

    some three times the amount of nitrogen n the soil beneathan overhead

    discharge

    than

    in

    the

    soil

    in

    the control

    plots,

    but

    unfortunately

    mea-

    surementswere taken

    only

    after the

    crops

    had

    been harvested. However

    Blackman

    1924)

    reported

    no

    appreciable

    differences

    n

    soil

    nitrogen

    con-

    tent before and after the

    application

    of electric fields to

    oats.

    A

    further

    unconfirmed

    uggestion

    by

    Priestley

    (1907)

    is

    that

    plants

    exposed

    to elec-

    trostatic

    ields

    may

    utilize

    atmospheric

    nitrogen

    directly,

    perhapsby

    com-

    bination of

    gaseousnitrogen

    with

    carbohydrates

    within

    the

    plant.

    Hart and

    Schottenfeld

    (1979)

    also observed

    that leaves

    of

    pole

    beans

    (Phaseolus

    multiflorus)

    ecame darker

    greenwhen

    exposed to

    electrostatic

    fields

    resulting

    in

    corona

    current

    from a

    few

    points on the leaf

    edges.

    Prolonged

    exposure

    caused loss of

    turgorand

    collapse

    of

    the plants.

    The

    pole

    bean

    plantswere

    grown

    in

    individual

    containers

    (20?C;

    30%

    relative

    humidity).Soil

    moisture

    content was

    monitored

    by

    measuring he

    resis-

    tance

    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 Hartand

    Schot-

    tenfield, 1979), and the mesh

    electrode

    was about 5 cm

    above the

    top of

    the plant. With the electrodechargednegatively, plants sustained2 ,uA

    of

    corona

    currentfor 8 hours

    with no

    visible

    effects,and with

    20

    AA

    of

    corona current

    here was

    little

    change

    in

    soil

    resistanceafter 5

    hours

    but

    the

    plant

    began to

    droop and one

    leaf

    became dark

    green and

    lost "tex-

    ture."

    At 50

    ,A,

    however,

    leaf

    discoloration

    developed

    rapidly and

    stem

    collapse

    occurred

    within 45

    minutes. With

    theelectrode

    charged

    positively

    and 20

    ,A

    of

    corona

    current,wilting

    occurred

    morerapidly

    and there

    was

    a

    large

    ncrease

    n

    soil

    resistance about

    35%

    n

    3

    hours).If

    slightly

    droop-

    ing plantswerewaterednear their stems they becameturgidagain.Hart

    and

    Schottenfeld

    attributed he

    effects

    o

    severe water

    oss from

    the leaves

    caused

    by the

    corona

    current.

    Bachman

    and

    Reichmanis (1

    973a) applied

    various

    strengthsof

    electric

    field to

    singleleaves of

    barley that

    were

    five days old

    and 5 cm

    longwith

    the cut end of

    the leaf

    in

    contact withthe

    negative

    electrode,and

    a

    positive

    electrode

    suspendedabove the

    leaf. Their

    results

    ndicated that

    with fields

    less

    than 40

    kV

    m-

    I

    leaf tip

    burning

    would not

    occur,butat a

    field

    strength

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  • 7/26/2019 Microwave Radiation and Plants

    13/54

    182

    THE BOTANICAL

    REVIEW

    of 400

    kV

    m-

    damage

    was

    almost

    instantaneous

    afteronly

    a few

    seconds

    of

    exposure).

    Above 800 kV

    m-

    '

    the

    density

    of

    positive

    air ions and

    level

    of ozone wereshownto buildup very rapidly.The time of the appearance

    of

    damage

    was

    found

    by

    Bachmanand

    Reichmanis

    to be

    proportional

    o

    1

    V2. They

    also investigated

    he

    speed

    of

    propagation

    of the

    damage

    down

    the leaf and found this

    to

    depend

    on the field

    strength.Although

    the

    damage

    at the

    tip appeared

    very quickly,

    the

    subsequentspread

    was at

    a

    reducedrate,e.g. for 250

    kV m-I

    damage

    appeared

    after

    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.

    Damage

    n

    air

    was

    compared

    o

    that

    in

    nitrogen

    and

    hydrogen

    by passing

    these

    gases

    over the leaf

    during

    electrification,

    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'

    exposure

    with 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

    damage

    was

    probablycausedby glowand brush

    dischargesat the leaf-

    air

    interface,

    so

    they

    made

    subsequent

    observations

    using

    a

    microscope

    in

    a dark

    room. They observed an

    ivory-colored glow

    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

    undamaged tissue.

    These

    observations,

    in

    their

    view, confirmedthat damage

    was caused by

    glow discharges.This

    damage

    mechanism is substantially

    different rom

    that earlier

    proposed by Murr

    (1964b) of

    acceleratedrespirationcausing

    membrane ruptureand cell dehydration.The results also differin other

    respects. Murr

    (1963a,

    1963b) reporteddamage at 40 kV

    m-l and below

    and an

    intermediatezone of

    dark

    green,

    while

    Bachmanand

    Reichmanis

    stated that no

    damage

    occurredbelow

    40

    kV

    m'-I

    and

    made no

    mention

    of color

    changes.

    After observingthe physical

    characteristicsof

    the damage, Bachman

    and Reichmanis

    (1973b)

    concentratedon the

    growth

    rates

    of plantssub-

    jected

    to

    electrostatic

    fields.

    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 strength

    corresponded o

    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 strong

    effects. By extrapolating hese

    growth rate

    results they pre-

    dicted that a

    zero value

    should occur at

    approximately800

    kV m-

    I

    which

    agreed

    with

    their earlier

    findings.

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  • 7/26/2019 Microwave Radiation and Plants

    14/54

    ELECTROSTATIC FIELDS, MICROWAVE RADIATION

    183

    Bankoske

    et al.

    (1976)

    investigated

    the effectsof 60

    Hz

    electromagnetic

    fieldsupon

    plants

    under

    power

    transmission ines.

    Theycarefullydesigned

    theirlaboratory xposurechamber o have an even field distributionwhen

    empty

    and then calculatedand measured he effectof

    placingplants

    nside.

    The field

    pattern was

    disturbed

    considerably

    and enhanced

    in

    the

    im-

    mediate

    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 considerable dis-

    tortion

    but

    if

    the

    plant height

    was

    less

    than 25% of the

    electrode

    spacing

    no more field enhancement occurredthan for a single plant beneath a

    transmission

    line.

    Plants

    with

    sharp-pointed

    eaves have a

    greater

    con-

    centrationof electric

    field

    aroundtheir

    tips

    and

    consequently

    breakdown

    owing

    to corona

    currentat lower

    voltages

    than broad-leaved

    plants.

    To define experimentalconditions Bankoskeet al.

    give "undisturbed"

    field

    strengths,

    .e. the

    field

    strength

    with an

    empty

    chamber,

    which bears

    no direct relation to the actual

    field strengths

    at the surfacesof

    leaves

    in

    the

    chamber.

    This is also

    recognized by

    Hart and

    Schottenfeld

    (1979)

    who

    preferred o state electricalexposure conditions

    in

    terms of

    AA

    of

    corona

    currentrather

    than

    unrepresentative

    ield

    strength

    values.

    Murr

    (1965a)

    uses

    a

    very simple

    model of

    a column

    of

    dielectric slabs

    to

    represent

    he

    air, leaves,

    roots and soil between the

    electrodes.This is

    unrealistic

    except

    in

    the one

    previously

    mentioned case of a

    very

    dense

    growthof

    plants whose top surface forms a new

    ground plane. Most of

    Murr's

    experimentsare

    with

    a

    single plant

    or

    groups of separatedplants

    (e.g. Fig. 5, Murr, 1965a)

    and so his values for

    field

    strengths are not

    valid,

    especially his dynamicfield strengthvalues.

    Enhancementof field strengthowing to the radiusofcurvatureof objects

    placed

    in

    electric fields means

    that

    it is

    impossible to

    relate the responses

    of

    different

    species

    of

    plants,

    or

    even different

    specimens

    of one

    species,

    to

    particular

    values

    of electrostaticfield

    strength.Each

    plant and part of

    a

    plant

    will

    experience

    different

    electrical forces at the

    leaf surface

    when

    placed

    in

    a

    uniformelectric

    field.Comparisonbetween

    experimentsbased

    on field

    strength

    values is

    extremely difficult.

    Bankoske

    et al.

    (1976) determinedthat corona

    currentwas responsible

    for leaf tip burningof corn(Zea mays) and alfalfa(Medicagosativa). The

    alternating

    field

    (60 Hz)

    had an

    undisturbed value of

    50

    kV

    m-1 and

    plants

    introduced

    into this had

    their uppermost leaf

    tips damaged after

    exposure

    for seven

    days.Corona

    dischargeswere

    photographed t the tips

    of

    leaves when

    plants

    were

    placed

    in

    "undisturbed"ield strengthsof 25-

    50

    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" ield

    strengths

    of 50

    kV

    m-l, 60

    Hz.

    The

    damage did not

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    184

    THE

    BOTANICAL REVIEW

    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 flutter

    owing

    to corona-inducedmotion.

    Murr

    1965b, 1966a,

    1966b, 1966c) nvestigated

    he

    effectof

    "reversed"

    (i.e.

    upperelectrodenegative)electrostatic ields,

    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

    flowed

    and caused damage

    to leaf

    surfaces.

    The

    thresholdvalues

    per plant

    fordamagewere 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

    epidermal

    damage

    and were believed to

    result

    in

    respiratory

    action suffi-

    cient

    to

    stimulate

    growth

    but

    not to cause substantial eaf tissue

    damage.

    Raising

    electric

    field strengths was

    thought

    to

    cause

    over-stimulation,

    enzyme

    toxicity

    and sufficient

    epidermaldamage

    to

    result

    n

    death of leaf

    tissue. Murr

    (1966a)

    concluded that currentbelow 10-16

    A

    per plant

    had

    no

    effect but growth stimulation occurred

    in

    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

    destruction occurs at

    10-5

    A

    and above.

    Blackman and Legg (1924), while examining

    the stimulation of

    the

    growth

    of

    barley by electric fields, used currents

    from 0.3

    x

    10-9 A

    to

    175 x 10-9 A per plant to maximize the response. The highercurrents,

    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) reported

    hat

    attempts

    to

    modify growth

    by using

    electro-

    static fields

    wereinconclusive, most having no

    effect or retarding

    growth.

    He

    also noted that

    lethaldamagemay be caused by

    coronadischarge

    rom

    leaf tips.

    Krueger

    and

    others (Kotakaand

    Krueger,

    1968; Krueger

    et

    al.,

    1978)

    have

    experimented

    with

    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

    increasing

    the

    air

    ion

    density (from

    the

    normal values of

    approxi-

    mately

    4.0

    x

    103 ions

    cm-3

    to

    approximately

    3.5

    x

    107

    ions

    cm-3)

    re-

    sulted

    n

    speededup respiration

    and

    growth

    ratesof

    young barley

    seedlings

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    ELECTROSTATICIELDS,

    MICROWAVE

    RADIATION 185

    (Kotaka

    et

    al.,

    1965)

    as

    compared

    to controls. Increases n air ion

    density

    have also been

    reported

    to increase

    the

    chlorophyll

    content

    slightly

    and

    the cytochrome content considerablyin treated plants (Kruegeret al.,

    1963), to speed

    up

    chlorosis

    of

    plants grown

    in

    iron-depleted

    environ-

    ments (Kotakaand

    Krueger,1968; Krueger

    t

    al., 1964),

    and to stimulate

    ATP

    metabolism

    (Kotaka

    et

    al., 1968)

    in

    isolated

    spinach chloroplasts.

    Complementing

    these

    results, Krueger

    et al.

    (1965)

    found that

    barley

    seedlingsgrown

    n ion-free

    atmospheres

    how retarded

    rowth,

    ack

    rigidi-

    ty

    and have soft leaves. Blackmanand

    Legg

    (1924)

    and

    Hicks

    (1957)

    have

    experimented

    with

    plants

    and

    trees

    in

    areas surrounded

    by

    wire

    mesh.

    All reportedretardedgrowth,lack of turgorand soft leaves. A wire mesh

    surrounding lants

    would act like a

    Faraday

    cage

    so the

    atmosphere

    nside

    it would be

    relatively

    free of electric fields and air ions.

    Krueger

    et al.

    (1978) exposed

    barley seedlings (Hordeumvulgare

    var.

    CaliforniaMari-

    out)

    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;

    current

    averaged

    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 latter and control

    groupsgrown

    with

    an ion-free

    and

    no

    electric

    field

    regime.

    They

    concluded that the air ions

    werebiologically active and responsible or the

    increases

    n

    plant growth,

    although they

    were unable to

    distinguish

    whether the effects were

    due

    to

    the air ions

    alone

    or to

    their

    presenceallowing

    electric currentsto flow

    through

    the

    plant. They pointed

    out that

    many previous

    workers

    had

    examined the

    responses

    of

    plants

    when

    subjected

    o electric

    fields but had

    not taken into

    account the presence

    and

    role of air

    ions.

    Bachman et al.

    (1971) found that under

    their experimentalconditions

    ozone appearedwith currentsof 5 MA hrougha singleleaf or 0.2 PA per

    leaf

    if

    therewere 20 plants. They recognized

    that the electric field at the

    tip

    of

    a

    barley

    leaf

    would be much

    higher

    than that calculated from the

    separation

    of leaf and

    electrode

    owing

    to the small

    radius of curvatureof

    the leaf

    tip.

    Also the

    edge

    of a

    barley

    leaf is covered with

    tiny "spikes"

    whose diameter s

    about 1000 times smaller

    than the leaf tip radius.Thus

    very high electric fields would exist around

    the "spike" tips and corona

    discharges

    will

    appear

    at much lower

    potentials

    than would be needed for

    discharges rom the leaf tip (equivalentto a few hundredV m'-Ibetween

    planarelectrodes),so ozone

    will

    be produced

    and lethal dischargecurrents

    can

    flow

    at

    quite

    moderate

    exposure levels.

    Ozone can

    damage plants

    and is

    produced at ground level by point

    dischargesduringthunderstorms.However,

    Shlanta and Moore (1972)

    commenting

    on

    the

    leaf

    tip burning

    of

    grassconsiderthatit is the discharge

    that

    burns he

    grassrather han the

    ozone.AlthoughBachmanet al. believe

    that

    it

    may

    inhibit

    growth

    and

    Menser et al.

    (1963) report ozone damage

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    186

    THE

    BOTANICAL

    REVIEW

    to

    tobacco

    leaves,

    complete

    death of a

    plant

    from

    the

    gas

    has not

    been

    suggested.

    Bankoskeet

    al.

    (1976)

    measured he amounts

    of ozone

    present

    when leaf tip burningoccurredduringcoronadischarge rom leaves. The

    level was

    16

    parts per

    billion

    (ppb) compared

    to an ambient level

    of

    8

    ppb.

    It was concluded hat ozone was

    unlikely

    to

    have caused

    any damage.

    Blackman

    et al.

    (1923) passed

    small currents

    0.5

    x

    10-10

    A

    per plant)

    through

    barley seedlings

    (Hordeumvulgare

    var.

    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 polarityproduceddefiniteincreasesin

    growth rate

    over controls (no

    electric field

    or current)which

    increased

    withtime and

    persisted or

    up

    to four hours

    after

    the

    currentwas

    stopped.

    The negative

    polarityresulted

    n

    an initial

    increasefollowed

    by

    a

    steadily

    decreasing

    rate

    of

    growth

    compared

    with 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

    charged

    set. Further

    work led

    them

    to concludethatneither he

    "electricwind" nor

    the

    gaseous

    byproducts i.e. ozone

    and

    nitrogenoxides)

    of the

    corona

    dischargewere

    responsible

    or the

    effecton the

    growth

    of the

    plants

    when a

    current

    passed.

    III.D

    CONCLUSIONS

    III.D

    (i) Summary

    Considerable

    effort was

    expended

    in

    the first third of

    this

    century

    in-

    vestigating

    the effects of

    electricfields on

    the growth

    of plants(Jorgensen

    and

    Priestley, 1914;

    Lemstrom,

    1904; Newman,

    1911).

    Although some

    of theresultswereveryencouraging e.g.fieldtrialswith cerealsand clover

    hay

    (Blackman, 1924)

    showed

    increases

    in

    yield of

    over 30%

    in half of

    the

    experimentsand

    yield increases

    n

    another

    28%of them], other

    work-

    ers

    could measureno

    effectsat all

    (Briggs

    t

    al., 1926;Collins et

    al., 1929).

    For

    18 years

    Lemstrom

    (1904) studied

    the

    effects of

    exposureto electric

    fields

    and

    reported

    healthy plants and

    increasedyields

    (includingcereals,

    vegetables

    and root

    crops). The time of day

    when

    electricitywas applied

    was

    importantsince

    exposureat

    midday or

    in hot sunshine

    could decrease

    crop yields. However, a reportof a subcommittee of the Board of Agri-

    culture

    and

    Fisheries

    (Committee of

    Electroculture,Final

    report of work

    carriedout

    1918-1937)

    indicated

    that, after

    prolonged

    research,benefits

    of

    electrical

    treatmentscould

    not be

    confidently

    predicted.

    Recently

    Kruegeret al. (1978)

    have

    shown that the density

    and type of

    air

    ions

    are

    important

    for

    plant growth,

    and

    suggested that ions rather

    than electric

    fields

    produced

    the

    various

    effects.

    Growth and respiration

    rates (Kotaka et

    al.,

    1965) as well as

    ATP

    metabolism (Kotaka et al.,

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    ELECTROSTATIC

    FIELDS,

    MICROWAVE

    RADIATION 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 burn-

    ing, 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 weedcontrol by high electric

    ields

    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

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    188

    THE

    BOTANICAL

    REVIEW

    whichwas

    sufficient

    o maintainthe leaf

    temperature

    t

    25?C.

    The

    authors

    calculatedthat a

    power

    level

    of 2

    mW cm-2

    was

    being

    absorbed

    by

    the

    leaf, and a minimum of 1 mW cm-2 represented he thresholdfor leaf

    protectionfrom frost at

    -

    5?C.

    After transfer o normal

    temperatures,

    he

    irradiatedplantremained

    healthy

    whereas the

    unprotected

    one had been

    destroyed

    by

    the cold.

    The

    investigators

    estimatedthat

    for frost

    protection

    125

    kW

    of

    power would

    be

    neededperhectarewith

    high capital

    cost but

    low

    running

    cost.

    In a

    later field

    test Bosisio et al.

    (1970)

    irradiated

    our-month-old

    corn

    (Zea

    mays) in a

    plot 8

    x

    8

    m

    and

    protected

    50%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

    irradiating

    ntenna

    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.

    Plants

    directlyexposed

    to

    the

    prevailing

    winds

    were,

    however,killed,

    even

    with

    150 mW

    cm-2

    radiation

    intensity.

    Microwave reatmentwould

    be

    very

    expensive

    and

    pose

    health

    hazards.

    Although

    Bosisio et al.

    (1970)

    measured

    only

    1

    mW

    cm-2

    at 3

    m

    from

    the

    edge

    of

    the

    plot, they

    had

    suggested

    earlier

    (Bosisio and

    Barthakur,

    1969)

    that farming

    areas using

    microwave

    radiation

    would have to be

    kept clear

    of

    people

    duringirradiations.

    Distributing

    the

    energy

    equally

    over

    large

    areas

    would also be

    a

    problem.

    Boulangeret

    al.

    (1969) produced a

    design

    study of a

    microwave grain

    drying

    system.

    Reducing grainmoisture content

    from

    20%

    to

    15% ook

    less than

    15 minutes by microwave

    methods (12

    kW,

    2450 MHz), 30

    minutes by

    a

    high

    frequency

    (HF) dielectricsystem

    (13

    MHz

    at

    10

    kW)

    and 150

    minutes

    by conventional hot-air

    techniques.

    Some

    wheat

    insects

    and their larvae were also controlled (Triboliumconfusum,Sitophilus

    granariusand

    Cryptolesteserrugineus).

    Costs

    for the electronic

    system

    were

    estimated

    to

    be half

    those

    of

    conventional

    methods. Baking and

    milling

    tests

    showed

    little differences

    between

    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

    practical

    imit to the

    increase in

    drying

    speed

    since too

    rapid

    heating

    led

    to crackingof

    the

    kernel and

    swelling

    of the

    grain,

    with

    consequentreduction

    n

    market

    grade.At

    very

    highratesof dryingthey found thatvaryingthe air flowthrough he grain

    had

    little effect on

    drying

    rateswhereas

    Boulanger

    et al.

    (1969)report hat

    the

    drying

    of

    wheat

    depends

    upon

    the

    change

    n

    vaporpressure

    determined

    by

    the

    air flow.

    Microwave

    radiation has been

    used

    to

    kill

    insect

    pests.

    Hightower et

    al.

    (1974)

    employed

    2450

    MHz

    radiation

    to control

    powder

    post

    beetles

    in

    imported

    hardwoods,

    particularly

    ellow pine and fir.

    Heating

    rates of

    30?C

    min'-I

    were achieved

    and the

    beetles and larvae

    could be controlled

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    ELECTROSTATICIELDS,

    MICROWAVE

    RADIATION

    189

    when wood

    temperatures

    of

    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 comparable o kiln dryingmethods butconsiderably asterand

    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

    treating nsect

    pests

    in

    grain

    and

    found

    that, often,

    much lower

    frequencies

    of radiation are more

    suitable

    (e.g. 39

    MHz)

    because the dielectric

    properties

    of

    insects

    and

    grains

    differ

    more from one

    another

    at lower

    frequencies

    han

    at microwave

    frequen-

    cies 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

    (Medicago

    sativa)

    has been re-

    duced

    (Nelson,

    1976b)

    to

    levels

    of

    5-15%

    from

    40-60%

    following

    ex-

    posure

    to 39 MHz

    and 2450

    MHz

    radiation. The

    success was

    related to

    seed moisture

    content,

    the

    greatestreduction

    n

    hard

    seed

    being

    n

    samples

    with the

    lowest

    moisture

    content. The

    optimum

    temperature

    was

    found

    to

    be 75?C.

    The

    benefits of

    treatment asted for

    up

    to four

    yearsafter the

    application.

    The

    effect was believed to be

    associated

    with the

    increase of

    water

    sorption

    by the seeds.

    In

    studies on the seeds

    of trees

    (Kashyap

    and

    Lewis,

    1974), germinationhas been

    found to

    increase