Pest Management Science Pest Manag Sci 56:299±308 (2000)

Why glyphosate is a global herbicide: strengths,

weaknesses and prospects†
Alan D Baylis*
Zeneca Agrochemicals, Jealott’s Hill Research Station, Bracknell, Berkshire, RG42 6EY, UK

Abstract: After 25 years of sales, glyphosate (N-(phosphonomethyl)glycine) is the world's biggest

selling and fastest growing agrochemical. Glyphosate has many strengths, particularly its systemicity
and ability to control perennial weeds, which have facilitated its role as an essential tool in modern
agriculture. Surveys of recent scienti®c and patent literature point to several key areas of current and
future interest in glyphosate and its use. Biotechnology topics have been the most popular, but there is
still much interest in the molecule per se, particularly in manufacturing processes. The mode of action
and physiology of glyphosate are common subjects. More applied aspects concern the development of
formulations with new adjuvants, or mixtures with other herbicides, to enhance performance. An
increasing number of papers are reporting on its environmental fate and on agronomic techniques
designed to improve the ef®ciency of application and minimise its environmental impact. Lack of weed
resistance is a key feature of glyphosate despite its widespread use. However, differential levels of
tolerance between species have led to changes in weed succession. There is a need to develop integrated
weed management systems in which glyphosate still plays a key role. Glyphosate is an exceedingly
effective herbicide and presents a considerable challenge to the development of new herbicides for
sustainable cropping systems and situations in which cost-effective and environmentally considerate
weed control is required.
# 2000 Society of Chemical Industry

Keywords: glyphosate; weed control; formulations; literature; patents, review

1 A GLOBAL HERBICIDE
Glyphosate (N-(phosphonomethyl)glycine) is a global
herbicide because of its versatility in controlling
economically a very broad spectrum of weeds under
varied agricultural, industrial, amenity and domestic
situations.
Since the introduction of the ®rst glyphosate
herbicide Roundup2 by Monsanto in the early
1970s, this chemical, in its several salts and many
formulations, has transformed global agriculture.
Together with other non-selective broad-spectrum
herbicides, particularly paraquat, it has freed millions
of people from the drudgery of manual weed control. It
has vastly diminished the constraints on productivity
imposed by perennial weeds. By any measure it has
become the world's leading agrochemical, and is still
the fastest growing,1 enabling the development of
novel and sustainable agricultural systems and facil-
itating the production of cheap and wholesome food.
Glyphosate has many strengths, and, as shown by its
outstanding record of success, few serious weaknesses.
It controls a very broad spectrum of weeds. Its
systemicity means excellent control of perennial
weeds, and its particular mode of action has meant
that any evolved resistance is exceedingly rare, unlike
that now facing many other widely used classes of
herbicide. It is of low toxicity and poses no risk of
leaching to ground-water. The more usually noted
weaknesses are: the need for higher rates to control the
generally more tolerant broad-leaved weeds; antagon-
ism by hard water and by many mixture partners; slow
speed of action and poor rainfastness compared to
paraquat. Glyphosate's lack of soil activity is a strength
considering crop safety, but a weakness concerning its
inability to control weeds emerging after application.
Overall it allows simple, cheap, ¯exible and effective
weed control while possessing excellent environmental
properties. These characteristics and bene®ts are
covered in earlier comprehensive reviews.2,3
The topics covered in this review re¯ect those of
current interest and are illustrated by reference to the
more recently published literature; some new data are
also included. The strengths and weakness of gly-
phosate will be considered, not only from the
perspective of the molecule as a herbicide, but also
from that of the global farming industry as a whole, ie

* Correspondence to: Alan D Baylis, Zeneca Agrochemicals, Jealott’s Hill Research Station, Bracknell, Berks RG42 6EY, UK
†
Based on a Paper presented at the meeting ‘Twenty-five years of increasing glyphosate use: the opportunities ahead’, organised by
J Caseley and LG Copping on behalf of the Crop Protection Group of the SCI and held on 23 February 1999 at the Royal Aeronautical
Society, London
(Received 30 April 1999; revised version received 15 October 1999; accepted 29 November 1999)

# 2000 Society of Chemical Industry. Pest Manag Sci 1526±498X/2000/$17.50 299

AD Baylis
its role as an essential tool in modern agriculture. In
this paper, all application rates are stated on a
glyphosate acid equivalent basis.
2 CURRENT RESEARCH INTERESTS
2.1 Scientific literature
A survey was conducted on glyphosate literature
published in scienti®c journals between mid-1995
and mid-1998 and abstracted in Chemical Abstracts,
Pestdoc from Derwent, or CABI (Centre for Agriculture
and Biosciences International) databases. (Table 1).
There were 630 papers in which some aspect of
glyphosate was a main topic. The following analysis
gives a contemporary view on the scope of research
into glyphosate and its use.
Biotechnology topics covering glyphosate-tolerant
(GT) crops and the use of glyphosate in GT crops,
were the most popular. There were 37 publications on
aspects of biotechnology per se, and 70 on weed control
in GT soybeans (Glycine max (L) Merr). Other crops
included cotton (Gossypium hirsutum L), oilseed rape
(Brassica napus L), maize (Zea mays L) and sugar beet
(Beta vulgaris L).
Papers concerned with conventional weed control
(ie non-transgenic crops) covered land preparation
(`burndown') before planting arable crops, control of
perennial weeds in crops prior to harvest (`pre-
harvest'), control of annual and perennial weeds in
temperate and tropical orchards, plantations, and non-
agricultural situations. A large number of publications
were concerned with glyphosate per se: its mode of
action, absorption, translocation and metabolism in
plants (especially with respect to environmental
in¯uences); its toxicology; analytical techniques; and
properties of various analogues.
The next largest category comprised the presenta-
tion of glyphosate in conjunction with particular
adjuvants or other herbicides, and the performance
of these formulations and mixtures. This was closely
followed by papers addressing ecological subjects, eg
the impact of glyphosate in the environment, its fate,
and agronomic techniques designed to minimise its
environmental impact. Application techniques cov-
ered conventional methods, and a particular issue is
that of spray drift. There were several papers on the
Table 1. Analysis of main topics covered in a survey of 630
papers on glyphosate published 1995–1998
Main topic No papers % Total
Glyphosate-tolerant crops 160
Conventional weed control 143
Properties of glyphosate 120
Formulations 76
Ecology 69
Application technology 29
Responses of weeds 17
Crop uses 16
300
use of `precision agriculture' to improve the ef®ciency
of glyphosate treatment.
The last two categories were concerned with effects
on plants: responses of weeds and weed populations,
and use of glyphosate on crops as a growth regulator.
Variations in susceptibility between species and
biotypes to glyphosate and changes in the importance
of particular species in weed ¯ora following repeated
use of glyphosate have been investigated. The crop
itself can be a target for glyphosate, with uses as a
desiccant prior to harvest, as a defoliation aid in
cotton, as a means of enhancing the sugar content of
cane and as a growth retardant all being reported.
2.2 Patent literature
A recent search of Chemical Abstracts revealed 696
patent publications pertaining to glyphosate. Twenty-
eight years after the publication of the ®rst patents
claiming the herbicidal properties of glyphosate, there
is clearly sustained, indeed increasing, commercial
interest in the utility of the molecule. One hundred
patent applications were published over the three years
prior to October 1998 (Table 2). The majority of
patents from this period cover claims to inventions in
the ®eld of glyphosate chemistry. These include
mainly manufacturing topics, but also esters and other
derivatives of glyphosate acid. A similarly large
number cover the genetic transformation of plants to
confer glyphosate tolerance. These patents cover not
only the biotechnology involved, but more often the
elite lines of crop plants created, and even mixtures of
glyphosate with other herbicides speci®cally for weed
control in transgenic crops.
Formulation patents include claims to novel types of
surfactant and other adjuvants, means of achieving
more concentrated formulations, and various presen-
tations, for instance as a granular product or even as a
seed dressing to control parasitic weeds. Effects
claimed usually involve better levels of weed control,
faster action or improved rainfastness, but also include
non-agronomic aspects such as lower irritancy.
There have been a number of mixture patents,
usually claiming synergy with other herbicides. Some
patents claim novel application methods and one
claims the use of glyphosate as a pharmaceutical
product.
Table 2. Analysis of main topics covered in a
survey of 100 patent publications on gly-
phosate published 1995–1998
Main topic No patents
25.4
22.7 Chemistry 34
19.0 Glyphosate-tolerant crops 29
12.1 Formulations 13
11.0 Mixtures 10
4.6 Formulation effects 9
2.7 Application techniques 4
2.5 Pharmaceutical uses 1
Pest Manag Sci 56:299±308 (2000)
 Why glyphosate is a global herbicide

These literature analyses illustrate current areas of
academic and industrial interest in glyphosate. Overall
they point to several key areas which will be discussed
in the remainder of this paper.
3 USE IN CROPPING SYSTEMS
3.1 Formulation
In recent years a vast array of different formulations of
glyphosate have been registered and sold. Being an
acid, the active ingredient in glyphosate products can
be formulated as salts or esters. This gives glyphosate
tremendous ¯exibility.
Glyphosate acid is generally less active than the
commercially available salts. This is believed to be a
result of poorer absorption into plants.4 At high rates,
particularly at low spray volumes, glyphosate acid can
precipitate out of solution. In the presence of cations
the solubility and physical form of glyphosate change.
The cations used in glyphosate herbicides greatly
increase solubility and make glyphosate more avail-
able. Absorption into leaves from dried spray deposits
is much higher and activity on plants is usually
substantially greater.
Results of a glasshouse experiment to compare the
activity of glyphosate acid to the trimethylsulfonium
salt (glyphosate-trimesium) are shown in Table 3.
Weeds (grasses tillering, broad-leaves with four to six
true leaves) were sprayed with a range of rates in
200 litre haÿ1 water (replicated three times). Applica-
tions were made with several primary C13/15 alcohol
surfactants (Synperonic series, ICI) varying in poly-
oxyethylene chain length5 and AL2042, an alkylpo-
lyglucoside used in the Touchdown2 formulation of
glyphosate-trimesium (Zeneca Agrochemicals). A
visual assessment of herbicidal activity (0±100%
control) was made 25 days after application and the
estimated rates for 90% control (ED90) were calcu-
lated from the rate response. Both acid and salt were
generally more effective with AL2042 or the longer
polyoxyethylene chain surfactants (A11 or A20), but
the salt was up to six times more active than the acid.
Formulation allows an active ingredient to be
presented to the target organism in an appropriate
way for the desired effect. By using appropriate cations
and adjuvants formulated at particular concentrations,
it has been possible to tailor glyphosate herbicides to
meet the needs of farmers, applicators and distributors
within a wide variety of agricultural and non-agricul-
tural systems. Some formulations have further im-
proved the environmental pro®le of products by using
surfactants with lower potential for irritancy and
reduced toxicity to aquatic organisms.
Liquid formulations are more common and suitable
for large-scale operations where ef®cient bulk-hand-
ling is required. In such situations, high concentrations
of glyphosate are preferred. On the other hand, low-
concentration liquid formulations have been devel-
oped which are appropriate for the needs of small-
holders who may wish to purchase small amounts of
glyphosate in a conveniently sized pack. A low
glyphosate concentration also facilitates the inclusion
of high concentrations of adjuvants which may
improve the degree and robustness of weed control.
Granular formulations have been introduced in some
markets. These enable higher concentrations of gly-
phosate and offer some advantages in operator
handling and easier pack disposal.
The choice of salt is an important consideration
when attempting to develop a particular type of
formulation. Although dimethylamine and mono-
sodium salts were mentioned in the ®rst publication
on the herbicidal properties of glyphosate,6 for many
years only the isopropylamine salt was sold. Currently,
in addition to the isopropylamine salt, trimesium,
ammonium and sodium salts are the basis of many
different formulations. These salts have been selected
for various reasons, eg compatibility with particular
adjuvants, herbicidal ef®cacy, cost and ease of
manufacture. Suitability for formulating as a liquid
or granular product is a good example of the

Table 3. Comparative activities of glyphosate acid and glyphosate-trimesium (G-TMS) applied with various non-ionic surfactants on annual grasses (G) and
broad-leaved weeds (B)

ED90(g haÿ1) a
Elymus repens Sorghum halepense Chenopodium album Abutilon theophrasti
Surfactant Glyphosate (G) (G) (B) (B)
Synperonic A2 Acid 953 f 1316 c 5197 c 3088 c
G-TMS 537 de 889 b 1091 b 1691 a
Synperonic A7 Acid 632 e 1334 c 3632 c 2869 c
G-TMS 500 d 753 b 1137 b 1961 b
Synperonic A11 Acid 476 cd 897 b 4000 c 2868 c
G-TMS 324 a 443 a 653 a 1535 ab
Synperonic A20 Acid 402 bc 734 b 3797 c 2385 c
G-TMS 366 ab 449 a 609 a 1553 a
AL2042 Acid 362 ab 761 b 3017 c 2706 c
G-TMS 383 b 445 a 510 a 1411 a
a
Means within each column followed by the same letters are not signi®cantly different at P = 0.05.

Pest Manag Sci 56:299±308 (2000) 301

AD Baylis
Figure 1. Equilibrium moisture contents of glyphosate mono-salts at 90%
(white bars) and 40% (black bars) relative humidity.
importance of the glyphosate salt. Salts of glyphosate
differ in hygroscopicity. Salts with high aqueous
solubility are generally too hygroscopic to allow the
manufacture of granular formulations.
A quantitative determination of hygroscopicity was
made by a gravimetric measurement of equilibrium
moisture contents of deposits of glyphosate salts.
Three drops of 10% acid equivalent solutions of
several mono-salts of glyphosate were applied to glass
slides and placed in a chamber where they could be
weighed having been allowed to equilibrate at 90% or
40% relative humidity. Duplicate measurements were
Figure 2. Deposits of glyphosate-trimesium and glyphosate-calcium. A mounted needle was drawn through the deposits from bottom to top in each case.
302
made. Equilibrium moisture contents decreased at
lower relative humidity, but some salts retained more
water (Fig 1). The least hygroscopic monosodium salt
has been used to make granular products. The ability
of spray deposits of glyphosate to retain and absorb
water after droplets dry down on leaf surfaces may be
important to uptake and consequent biological activ-
ity.4,7
Hygroscopicity and physical form of dried deposits
of glyphosate salts can be demonstrated by observing
how readily they re-hydrate. Figure 2 illustrates this by
showing the path of a mounted needle drawn through
deposits formed from drops of glyphosate salt solu-
tions, applied to glass slides, desiccated and then
returned to ambient conditions (21 °C, 40% relative
humidity). The highly water-soluble trimesium salt
(solubility 1050 g litreÿ1) will spontaneously re-hy-
drate, but less-soluble salts, eg calcium which has a
solubility of 0.3 g litreÿ1, form a glass-like deposit. In
fact, sub-optimal performance of glyphosate may be
associated with the presence of calcium and other
polyvalent ions on leaf surfaces.8 Very poor absorption
of glyphosate in the presence of calcium has been
recorded.9
3.2 Mode of action and symptomology
Glyphosate inhibits 5-enolpyruvylshikimate 3-phos-
phate synthase (EPSPS; E.C. 2.5.1.19) and the
literature on its mode of action has been thoroughly
reviewed.3 EPSPS is located primarily in plastids,
although a cytoplasmic form exists also. Inhibition of
Pest Manag Sci 56:299±308 (2000)

EPSPS results in an accumulation of shikimate-3-
phosphate, ultimately leading to a block in the
production of aromatic amino acids with resultant
effects on protein synthesis.
Recent work has surmised that glyphosate is not a
transition state inhibitor competing with phosphoe-
nolpyruvate (PEP), as previously believed, but that its
binding behaviour with EPSPS is better described as
an adventitious allosteric interaction, ie a signi®cant
part of the glyphosate molecule binds outside the
EPSPS active site, and that a conformational change
upon glyphosate binding makes the active site un-
available to PEP.10
Although EPSPS is the only known enzyme target of
glyphosate,11 it affects many physicochemical and
physiological processes.12 Among these are reductions
in photosynthesis and degradation of chlorophyll;
inhibition of auxin transport and enhancement of
auxin oxidation. These aspects of the mode of action
of glyphosate are either a direct consequence of the
blockage of the shikimate pathway (through which
some 30% of assimilated carbon is estimated to pass)13
or a result of feedback mechanisms. Metal chelation
may also be signi®cant in restricting the availability of
polyvalent cations, eg as enzyme co-factors or as key
structural or functional elements, eg Mg2‡ in chloro-
phyll.12
Glyphosate is regarded as a slow-acting herbicide,
and this has been seen as a weakness. However,
physiological effects in treated plants can be detected
very much earlier than the ®rst appearance of
herbicidal symptoms. Large increases in concentra-
tions of shikimic acid occurred within 24 h of gly-
phosate application to Betula papyrifera Marsh (white
birch) even from rates which showed no visible
symptoms.14 A diagnostic test for determining
whether any crop phytotoxicity related to spray drift
can be attributed to glyphosate involves an assay for
determining the concentration of shikimate in affected
plants.15 In white birch, sub-lethal doses of glyphosate
reduced photosynthesis, chlorophylls a and b declined,
and carotenoids increased.14 All these modes of action
result in the familiar symptoms of glyphosate phyto-
toxicity: chlorosis, pigmentation, stunting, and reduc-
tion in apical dominance.
A further symptom of glyphosate phytotoxicity
which occurs very rapidly in some species is an
epinastic response, which is typical of the effects of
ethylene. Amaranthus retro¯exus L is unusually suscep-
tible for a broad-leaved weed, and the effects of
glyphosate can be observed only a few hours after
spraying. Ethylene production in B papyrifera in-
creased following glyphosate application,14 but little
change in ethylene production was observed in gly-
phosate-treated sugar beet (B vulgaris).16
Although primarily a herbicide, of course, the
broader utility in the effect of glyphosate on plants is
better illustrated by its uses as a growth regulator.
Glyphosate can be used as a desiccant to facilitate the
harvesting of high-moisture-content crops, eg soy-
Pest Manag Sci 56:299±308 (2000)
Why glyphosate is a global herbicide
beans and oilseed rape, especially in situations of late
harvest or lodging. The symptoms of glyphosate action
in plants are not necessarily always herbicidal. Gly-
phosate and its analogue glyphosine have been used to
enhance sucrose concentrations in sugar cane at
harvest for many years.17 More recently, it was
reported that in Australia, 135 g haÿ1 glyphosate
helped to maintain productive pastures of bent grass
(Agrostis castellana Boiss & Reut) by supressing seed-
head development and reducing apical dominance
such that new tillers dominated the sward.18 This
increased digestible dry matter and protein content
compared with the unsprayed control which was
dominated by seedheads. Applications of glyphosate
before spraying cotton defoliants inhibited the re-
growth which is a common problem of cotton in wet
seasons when harvesting is delayed more than two or
three weeks after defoliation.19
3.3 Weed control
In any weed control situation the necessary rate is
determined by the most tolerant species. A range of
glyphosate rates are recommended for weed control
from a few hundred g haÿ1 to several kg haÿ1. Inherent
tolerance to glyphosate in plants can be considered
between and within species.
3.3.1 Between species: tolerance
Higher rates are usually required for perennial weed
control. These species are usually large plants with a
great potential for regeneration. However, amongst
annual weeds, relative potency can vary several-fold
(Table 3). Grasses are usually more susceptible to
glyphosate than broad-leaved weeds. However, there
can be surprisingly wide differences between closely
related species in their degree of tolerance. Clear
differences in the susceptibility of various species of
Amaranthus were recorded in a glasshouse experi-
ment. Weeds (six to eight true leaves) were sprayed
with a range of rates in 200 litre haÿ1 water (replicated
three times). A visual assessment of herbicidal activity
(0±100% control) was made 22 days after application.
Results illustrating differences in control from 225
g haÿ1 are shown in Table 4. While A retro¯exus, as
Table 4. Control of Amaranthus
spp by glyphosate-trimesium
(225 g haÿ1) 22 days after appli-
cation.
Control a
(%)
A retro¯exus 94 a
A albus 89 ab
A viridis 77 b
A tamariscinus 58 c
A hybridus 52 c
a
Means followed by the same
letters are not signi®cantly differ-
ent at P = 0.05.
303
AD Baylis
previously noted, is a broad-leaved weed very
sensitive to glyphosate, other species are more
tolerant. This may be signi®cant, in that common
waterhemp (A tamariscinus Nutt) has been noted as a
potentially increasingly serious problem in GT soya.20
However, this species tends to emerge later in the
season, after application of glyphosate, and a residual
herbicide may be needed for good season-long
control.
The reasons for tolerance of a particular species may
be due to many factors, possibly in combination.
Studies on the reasons for tolerance are rare. Calcium
is a well-known antagonistic ion when present in water
used for spraying.21 However, calcium and magne-
sium, which is similarly antagonistic, are also present
on the leaf surfaces of many plants.8 More information
on tolerance is available from studies on different
biotypes of the same species.
3.3.2 Within species: tolerance and resistance
A single population of ®eld bindweed (Convolvulus
arvensis L) was found to contain ®ve biotypes which
differed in their susceptibility to glyphosate.22 These
were clearly distinguishable on the basis of their
morphology. The two biotypes with extremes of
tolerance were selected for study. The isopropylamine
salt of glyphosate at 2.24 kg haÿ1 easily controlled the
more susceptible biotype, but the more tolerant
biotype was much less affected, increasingly so when
older plants were compared. No differences in gross
absorption and translocation of glyphosate in the two
biotypes were recorded.23 In suspension culture, cells
of the more susceptible biotype had higher initial rates
of glyphosate uptake.24 The EPSPS from both
biotypes was equally sensitive to glyphosate. However,
although both biotypes initially had similar levels of
enzyme activity at the time of treatment, it then
increased in tissues of the tolerant biotype. It was
postulated that this was a consequence, not a cause of
better survival by this biotype. There was greater ¯ow
of carbon through the shikimate pathway in the
tolerant biotype, a greater pool of phenylalanine and
overall greater growth rate. In conclusion, it seems that
multiple mechanisms are involved in the greater
tolerance of this biotype. Indeed, genetic analysis has
indicated the involvement of multiple genes.25
Many weed species have evolved resistance to
herbicides of many classes. These situations have
arisen following high selection pressures being exerted
by repeated use of a herbicide. This has encouraged
naturally occurring mutations which do not bear a
®tness penalty, or previously small pre-existing popu-
lations of more tolerant biotypes, to become more
signi®cant in the weed ¯ora. The International Survey
of Herbicide Resistant Weeds lists 212 biotypes of
weeds which are resistant to 15 classes of herbicide.26
Of these, there are 50 biotypes resistant to ALS
inhibitors, for example. However, only one, a Lolium
rigidum Gaud biotype from Australia, is resistant to
glyphosate.27 A further case involving Eleusine indica
304
(L) Gaertn in Malaysia has been reported in the papers
from this symposium.28
The lack of weed resistance to glyphosate has proved
to be unusual considering its widespread and intensive
use. Several factors may contribute to this:29 the poor
®tness of any genotypes evolving with an EPSPS
enzyme resistant to glyphosate, ie a mutated enzyme
which may not be so affected by glyphosate but which
is also less ef®cient at processing PEP; the absence of
ef®cient metabolic degradation or deactivation of
glyphosate by plants; the lack of soil activity, with the
result that only the part of the gene pool present in
plants actually sprayed is selected upon; and the
unique mode of action and chemistry of glyphosate
among commercial herbicides (ie other herbicides
used for weed control in the same ®eld will not
contribute to the selection pressure).
In future, resistance problems may be posed by GT
crop volunteers and any new biotypes of weeds arising
through out-crossing and transfer of glyphosate
tolerance genes.29
3.4 Integrated weed management
Although resistance to glyphosate is extremely rare,
clearly species and even biotypes differ in their
response. This varying tolerance between plants gives
rise to the very practical problem of weed succession.
With the advent of GT crops, there has been much
speculation on the extent to which the importance of
particular species in the weed ¯ora in crops will change
and the speed at which this will occur. In plantations in
SE Asia where glyphosate has been used very
intensively, there has been a marked rise in the
dominance of annual broad-leaved weeds, eg Asystasia
spp, Heydyotis spp, Borreria spp. Several spray rounds
each year are required to achieve acceptable weed
control. Programmes have been introduced in which
integrated spray rounds of paraquat and glyphosate-
trimesium will give total vegetation control without
further encouraging weed succession.30 Increasing
consideration is also given to the impact of particular
weed species. For example, the presence of some
grasses is tolerated, as these are less competitive and
their root systems help to prevent soil erosion.
Euphorbia heterophylla L is a major weed of soybeans
in Brazil which has become resistant to formerly widely
used ALS inhibitors, such as imazethapyr. Weed
pressure from this species can be particularly high
because of its emergence over a prolonged period
before and after drilling and re-growth from adventi-
tious buds after application of contact herbicides. A
weed management system has been devised which
involves controlling weeds 20 days before drilling with
glyphosate-trimesium, followed by paraquat immedi-
ately before drilling, and then application of the
selective herbicide fomesafen three to ®ve weeks after
drilling.31
An alternative approach integrates weed control
using cultivation and glyphosate.32 Weed germination
prior to drilling winter crops was encouraged by tillage
Pest Manag Sci 56:299±308 (2000)

and weeds were subsequently controlled by gly-
phosate-isopropylammonium. This reduced autumn
weed populations facilitating effective and economic
weed control by selective herbicides. Overall herbicide
use over ®ve years was reduced by more than 50%.
Although many mixtures with selective herbicides
were antagonistic to glyphosate, overall they improved
post-emergence broad-leaved weed control in GT
soya.33 Another bene®t of mixtures with soil residual
activity is to provide protection against weeds germi-
nating after spraying. Mixtures of herbicides with
different modes of action may help forestall the
development of weed resistance.
Low rates of glyphosate were preferred to higher
rates in order to maintain a balanced weed population
in almond orchards.34 This improved water in®ltra-
tion, provided a habitat for bene®cial insects and
helped to preclude invasion by aggressive weeds more
tolerant of glyphosate, such as sedges.
3.5 Application
3.5.1 Spray drift
The use of hydraulic nozzles results in the formation of
spray droplets with a wide size distribution. Smaller
droplets are more likely to be retained on leaf surfaces,
but those with diameters of < 100 mm are generally
considered to be susceptible to drift. Usually, any
glyphosate spray drift reaching crops will be at
relatively low rates. This, and the slow development
of symptoms, may mean that phytotoxicity is not
necessarily obvious. However, care has to be taken as
deleterious effects can occur. Herbicides were applied
to vines in the autumn at low rates simulating drift. In
the following spring glyphosate, especially when mixed
with 2,4-D, affected growth rate of the vines.35
Glyphosate caused fruit drop of citrus, especially if
the fruit itself was contacted by spray.36 Fully shielded
booms caused less fruit drop than open-boomed
sprayers.
The advent of GT soybeans might potentially
increase the risk of glyphosate drift causing phytotoxi-
city to young maize.37 Maize injury from glyphosate
spray drift was greater when GT soya was sprayed at
lower water carrier volumes.38
Some drift-management adjuvants are available.
Reductions in the number of drift-prone droplets
resulted in less drift in a wind-tunnel experiment.39
Surfactants themselves vary in their effects on droplet
size distribution. At the usual use concentrations, a
nonylphenyl surfactant with eight ethylene oxide units
produced a lower proportion of driftable droplets than
a tallow amine surfactant with 15 ethylene oxide
units.40 However, mechanical factors such as nozzle
type and water volume produce the greatest variation
in droplet sizes. Data on spray characteristics of a
formulation of glyphosate-trimesium are shown in
Table 5. The formulation was sprayed at several
concentrations from a pressurised container using a
single nozzle. Different nozzles and pressures were
used to achieve three different ¯ow rates. At a forward
Pest Manag Sci 56:299±308 (2000)
Why glyphosate is a global herbicide
speed of 8 km hÿ1 these correspond to application
volumes of 100, 200 and 400 litres haÿ1. Droplet sizes
were measured using a Malvern 2600 Particle Sizer
®tted with a 1000 mm objective lens. The volume
median droplet diameter (50% of spray volume is
found in droplets smaller or larger than this diameter)
decreased and the proportion of spray volume in
potentially driftable droplets increased under the
conditions simulating lower volume applications and
higher rates of formulation, which would give a higher
concentration of surfactant.
3.5.2 Precision agriculture
Over recent years, the availability of many new and
rapidly advancing technologies ± information technol-
ogy, satellites, imagery, etc ± have combined with ever
more sophisticated agricultural machinery to offer
farmers the means to tailor and apply crop inputs with
far greater accuracy and ¯exibility. `Precision agricul-
ture' is becoming more and more widely adopted,
especially in the corn/soya belt of the US Mid-West.
Typical applications at present are variable fertiliser
and residual herbicide rates according to soil maps (of
nutrient and organic matter status) using satellite
navigation. Precision application of post-emergence
herbicides is not widely practised at present, but would
be very attractive if its development had economic
bene®ts. At present, glyphosate is often used inef®-
ciently because weeds occur in patches, the suscept-
ibility of individual weed species in a mixed ¯ora can
vary several-fold, and environmental conditions at the
time of spraying can affect the degree of weed control
obtained from the rate applied.
Spraying patches demands that weeds can be
located in some way.41 While in some situations it
seems possible to map weed patches by infestations in
the previous season, it would obviously be preferable
to detect weeds shortly before spraying. The presence
or absence of a crop will in¯uence how easily this can
be achieved. Sprayers with weed-sensing systems have
been available in N America for a number of years.42
The presence of weeds may be detected by determin-
ing the differential re¯ectance of red and near-infra-
red wavelengths between green tissue and soil.
Forward-mounted sensors signalled the operation of
each nozzle independently as weeds were detected in
the spray path. Using such a sensor system allowed the
rate of application of a mixture of glyphosate and
dicamba to be reduced by up to 60% over three years,
and gave comparable results to conventional spraying
on 80% of occasions.43 Best results were obtained
when weed occurrence was patchy, but it was observed
that small weeds, particularly grasses, at low popula-
tion densities were dif®cult to detect successfully.44
When spraying weeds in stubble, detection was more
dif®cult in taller stubble. Detection was sub-optimal at
low sun angles. Besides inaccurate detection, any
requirements to drive spraying equipment more slowly
may limit the advantages of this sort of system.
The ability to distinguish between species differing
305
AD Baylis
Flow rate
Nozzle (litre minÿ1)
b
Fine 0.60
Mediumc 1.31
Coarsed 2.61
a
VMD = volume median diameter (mm), DD = %spray volume in droplets < 100 mm diameter.
b
Table 5. Effect of spray concentration of product Lurmark 015-F110 with spray at 3 bar pressure.
c
and nozzle aperture on spray characteristicsa of Lurmark 04-F110 with spray at 2 bar pressure.
d
Touchdown2 Lurmark 08-F110 with spray at 2 bar pressure.
in susceptibility would be a further improvement
reducing overall application rates. Various imaging
techniques involving plant morphology, leaf colour
and shape are being researched.45,46 `Smart' spraying
systems for glyphosate would obviously have to offer
economic advantages, but there may be environmental
pressures to adopt precision application. More accu-
rate application could also bene®t farmers directly, of
course, by reducing phytotoxicity from spray drift.
In the future, decision support software could ®ne-
tune glyphosate rates according to prevailing and
forecast environmental conditions. Environmental
factors are very signi®cant to glyphosate activity.47
These can have effects both on the plant, conditioning
its susceptibility, and on the herbicide directly. For
example, although 820 g haÿ1 glyphosate-isopropyla-
mine gave good control of Chenopodium album L and
Abutilon theophrasti Medik in GT soya in 1994, control
in 1995 was poor.33 In 1995, weeds were at the same
growth stage, but markedly smaller. 1995 had been
drier and relative humidity was lower at spraying,
although air temperatures were similar. Glyphosate is
less effective in controlling drought-stressed weeds.48
It was also less effective at lower temperatures on
plants grown where soil water holding was maintained
at ®eld capacity. Glyphosate absorption and transloca-
tion is often impaired in droughted plants and those
grown at low temperatures.49 Under drought stress,
however, reduced glyphosate ef®cacy was associated
with higher temperatures.48 Increased evapo-tran-
spiration may have exacerbated the effects of soil
moisture de®cit. Glyphosate was more effective at
higher relative humidities, and this may be because of
greater absorption.50 Leaves of C arvensis had less
cuticular material when grown at higher relative
humidity, possibly facilitating better absorption.23
4 PROSPECTS
Over the past 25 years many new herbicides have been
developed and sold. Many have been far more potent,
most have been faster acting or had other advantages,
but none have had such a potential to contribute to
global weed control in such a thorough, versatile and
sustainable way as glyphosate.
As the world's most widely used and fastest growing
pesticide there is understandable interest in the impact
306
Touchdown
0% 1.25% 2.50% 5.00%
VMD a
DD a
VMD DD VMD DD VMD DD
146 31 132 36 116 41 112 43
284 9 247 13 222 17 223 19
423 5 356 6 312 19 264 14
glyphosate has on the environment. The mode of
action of glyphosate suggests that toxicity would be
con®ned to life forms possessing the shikimate path-
way, ie plants and micro-organisms. Extensive re-
search has demonstrated that glyphosate has, indeed,
very low toxicity to mammals, birds and ®sh; it does
not bioaccumulate; glyphosate and its metabolites are
not harmful to soil fauna and micro¯ora or honey-
bees.3
Glyphosate is rapidly adsorbed in soil, making it
essentially immobile. Soil binding is reversible and on
desorption glyphosate is degraded by various bacteria
to aminomethylphosphonic acid (AMPA) or sarco-
sine, and ultimately to inorganic phosphate, ammonia
and carbon dioxide.3
Environmental issues surrounding the introduction
of crops genetically modi®ed (GM) to be tolerant to
glyphosate or other herbicides are of much current
interest and the impact of these has already been
reviewed.51 Although glyphosate itself is particularly
benign, it indirectly in¯uences the environment by the
consequences of its use in agriculture. For example, by
chemically controlling weeds it enables `conservation
tillage' systems to be adopted by farmers direct drilling
into uncultivated ground, saving fuel, preventing soil
erosion and improving soil structure.52 In GM tolerant
crops, controlling weeds post-emergence with gly-
phosate not only avoids the prophylactic use of pre-
emergence soil residual herbicides, but allows the
development of a weed ¯ora to support birds and
bene®cial insects. Unsprayed cereal headlands in the
UK had a greater number of arthropod species
important in the diet of young birds of many species,53
and a similar situation might be expected in GM
tolerant sugar beet early in the season before control-
ling weeds with glyphosate. Weeds do not always need
to be removed completely and, if suppressed, the cover
remaining will preclude invasion by more noxious
species. Such systems involving glyphosate pro-
grammes have also provided a habitat for bene®cial
insects.34
This paper has covered the key aspects of gly-
phosate's role in modern weed control. What are the
prospects of superior herbicides being discovered and
commercialised?
Clearly, it is possible to note individual weak points
in the technical pro®le of glyphosate, eg it is less
Pest Manag Sci 56:299±308 (2000)

effective on broad-leaved weeds and lacks soil residual
activity. Yet, there are always counterpoints, eg the
extreme rarity of weed resistance and absence of carry-
over on to crops. And there are always solutions if
needed, eg mix with other herbicides.
Glyphosate has not only been a means of weed
control, but it has shaped new agricultural systems
around it, most notably glyphosate-tolerant crops.
This now places a new hurdle in the path of developing
new and better herbicides. In addition to being
superior commercial weedkillers, they must be su-
perbly selective to many crops. If not inherently
selective, then not only must they be capable of being
used selectively by biotechnology, but then must also
rely on varieties expressing the resistance trait being
successfully commercialised. Other developing tech-
nologies in precision application may mean that the
overall ef®ciency and economy with which glyphosate
can be used will present considerable challenges to
even the most potent of potential new contenders.
ACKNOWLEDGEMENTS
I thank Cliff Hart, Dr Anne Waller, John Tobbut and
the Zeneca Agrochemicals Non-Selective Herbicides
team for data and Dr Ceris Jones and Sarah Harvey
Speck for assisting with the compilation of literature.
REFERENCES
1 Woodburn A, Glyphosate: Production, pricing and use world-
wide. Pest Manag Sci 56:309±312 (2000).
2 Grossbard E and Atkinson D (eds), The Herbicide Glyphosate,
Butterworths, London. 490 pp (1985).
3 Franz JE, Mao MK and Sikorski JA, Glyphosate: A unique global
herbicide. American Chemical Society Monograph 189, Wa-
shington DC.653 pp (1997).
4 MacIsaac SA, Paul RN and Devine MD, A scanning electron
microscope study of glyphosate deposits in relation to foliar
uptake. Pestic Sci 31:53±64 (1991).
5 Hollis GL (ed), Surfactants Europa, 3rd edn, The Royal Society of
Chemistry, Cambridge. 16 pp (1995).
6 Baird DD, Upchurch RP, Homesley WB and Franz JE,
Introduction of a new broad-spectrum post-emergence herbi-
cide class with utility for herbaceous perennial weed control.
Proc North Central Weed Control Conf 26: 64±68 (1971).
7 Nalewaja JD, Matysiak R and Freeman TP, Spray deposit
residuals of glyphosate in various carriers. Weed Sci 40:576±
589 (1992).
8 Hall GJ, Hart CA and Jones CA, Plants as sources of cations
antagonistic to glyphosate activity. Pest Manag Sci 56:351±358
(2000).
9 Nalewaja JD, Devilliers B and Matysiak R, Surfactant and salt
affect glyphosate retention and absorption. Weed Res 36:241±
247 (1996).
10 Sikorski JA and Gruys KJ, Understanding glyphosate's molecu-
lar mode of action with EPSP synthase: evidence favoring an
allosteric inhibitor model. Acc Chem Res 30:2±8 (1997).
11 Siehl DL, Inhibitors of EPSP synthase, glutamine synthase, and
histidine synthesis. Rev Toxicol 1:37±67 (1997).
12 Cole DJ, Mode of action of glyphosate ± a literature analysis, in
The Herbicide Glyphosate, ed by Grossbard E and Atkinson D,
Butterworths, London. pp 48±74 (1985).
13 Singh BK, Siehl DL and Connelly JA, Shikimate pathway: why
does it mean so much to so many? Oxford Surv. Plant Mol Cell
Biol 7:143±185 (1991).
Pest Manag Sci 56:299±308 (2000)
Why glyphosate is a global herbicide
14 Stasiak MA, Hofstra G and Fletcher RA, Physiological changes
induced in birch seedlings by sub-lethal applications of
glyphosate. Can J Forestry Res 22:812±817 (1992).
15 Singh BK and Shaner DL, Rapid determination of glyphosate
injury to plants and identi®cation of glyphosate-resistant
plants. Weed Technol 12:527±530 (1998).
16 Cole DJ, Caseley JC and Dodge AD, In¯uence of glyphosate on
selected plant processes. Weed Res 23:173±183 (1983).
17 Boehm RA, Increasing pro®tability with ripeners and specialist
adjuvants. Proc West Indian Sugar Technologists 26th Conference,
pp 278±279 (1997).
18 Hill RD, Missen DJ and Taylor RJ, Use of glyphosate to prevent
development of reproductive tillers and extend vegetative
growth of bent grass (Agrostis castellana). Aust J Exper Agric
36:661±664 (1996).
19 Martin-Duval T, Vargas R, Wright S and Roberts B, Pre-
conditioning cotton with glyphosate for defoliation. Proc
Beltwide Cotton Conf 2:1479±1480 (1997).
20 Owen MDK, North American developments in herbicide-
tolerant crops. Proc Brighton Crop Prot Conf ± Weeds, BCPC,
Farnham, Surrey, UK. pp 955±963 (1997).
21 Nalewaja JD and Matysiak R, Salt antagonism of glyphosate.
Weed Sci 39:622±628 (1991).
22 DeGennaro FP and Weller SC, Differential susceptibility of ®eld
bindweed (Convolvulus arvensis) biotypes to glyphosate. Weed
Sci 32:472±476 (1984).
23 Westwood JH, Yerkes CN, DeGennaro FP and Weller SC,
Absorption and translocation of glyphosate in tolerant and
susceptible biotypes of ®eld bindweed (Convolvulus arvensis).
Weed Sci 45:659±663 (1997).
24 Westwood JH and Weller SC, Cellular mechanisms in¯uence
differential glyphosate sensitivity in ®eld bindweed (Convolvu-
lus arvensis) biotypes. Weed Sci 45:2±11 (1997).
25 Duncan CN and Weller SC, Heritability of glyphosate suscept-
ibility among biotypes of ®eld bindweed. J Heredity 78:257±
260 (1987).
26 Heap I, International Survey of Herbicide Resistant Weeds [On-line]
Internet. Available: www.weedscience.com. [1998 December
3].
27 Powles SB, Lorraine-Colwill DF, Dellow JJ and Preston C,
Evolved resistance to glyphosate in rigid ryegrass (Lolium
rigidum) in Australia. Weed Sci 46:604±607 (1998).
28 Lee LJ and Ngim J, A ®rst report of glyphosate-resistant
goosegrass (Eleusine indica) in Malaysia. Pest Manag Sci
56:336±339 (2000).
29 Bradshaw LD, Padgette SR, Kimball SL and Wells BH,
Perspectives on glyphosate resistance. Weed Technol 11:189±
198 (1997).
30 Hee LC, Kim LJ and Jantan B, Comparative studies of a
paraquat mixture and glyphosate and/or its mixtures on weed
succession in plantation crops. The Planter, Kuala Lumpur
69:525±535 (1993).
31 Vidal RA, Fleck NG, Oliveira NA, Strello R, Guimaraes FB and
Silva NG, Increasing the number of mechanisms of action of
herbicides for management of weed resistance. Proc Brighton
Crop Prot Conf ± Weeds, BCPC, Farnham, Surrey, UK. pp 363±
368 (1997).
32 Hutcheon JA, Stride CD and Wright KJ, Manipulation of weed
seedbanks in reduced tillage systems for sustainable weed
control. Asp App Biol 51:249±254 (1998).
33 Lich JM, Renner KA and Penner D, Interaction of glyphosate
with post-emergence soybean (Glycine max) herbicides. Weed
Sci 45:12±21 (1997).
34 Neu JL and Sieckert EE, Weed suppression with low rates of
Roundup for orchard ¯oor management. Proc California Weed
Sci Soc 47:202 (1997).
35 Bhatti MA, Al-Khatib K and Parker R, Wine grape (Vitis vinifera)
response to fall exposure of simulated drift from selected
herbicides. Weed Technol 11:532±536 (1997).
36 Erickson CG, Management of glyphosate-related citrus fruit
drop. Proc Florida State Hort Soc., pp 40±42 (1996).
307
AD Baylis
37 Matthews JL and Kapusta G, Corn tolerance to simulated
glyphosate and glufosinate drift. Res Rep North Central Weed Sci
Soc 53:505±506 (1996).
38 Franzenburg DD, Owen MDK and Lux JF, Glyphosate drift on
corn as affected by carrier volume. Proc North Central Weed Sci
Soc 50:19±20 (1995).
39 Downer R, Wolf TM, Chapple AC, Hall FR and Hazen JL,
Characterizing the impact of drift management adjuvants on
the dose transfer process. Proc Fourth Int Symp Adjuvants for
Agrochemicals. pp 138±143 (1995).
40 Miller PCH, Butler Ellis MC and Tuck C, The in¯uence of
adjuvants on droplet production. Proc Fourth International
Symp Adjuvants for Agrochemicals pp 95±102 (1995).
41 Miller PCH, Stafford JV, Paice MER and Rew LJ, The patch
spraying of herbicides in arable crops. Proc Brighton Crop Prot
Conf ± Weeds, BCPC, Farnham, Surrey, UK. pp 1077±1086
(1955).
42 Felton WL, Commercial progress in spot spraying. Proc Brighton
Crop Prot Conf ± Weeds, BCPC, Farnham, Surrey, UK. pp
1087±1096 (1995).
43 Blackshaw RE, Molnar LJ, Chevalier DF and Lindwall CW,
Merits of a weed-sensing sprayer to control weeds in
conservation fallow and cropping systems. Weed Sci 46:120±
126 (1998).
44 Blackshaw RE, Molnar LJ, Chevalier DF and Lindwall CW,
Factors affecting the operation of the weed-sensing Detect-
spray system. Weed Sci 46:127±131 (1998).
45 Bull CR, Zwiggelar R and Stafford JV, Imaging: a technique for
assessment and control in the ®eld. Asp App Biol 43:197±204
(1995).
308
46 Woebbecke DM, Meyer GE, Bargen K and Mortensen DA,
Color indices for weed identi®cation under various soil,
residue and lighting conditions. Trans Am Soc Agric Eng
38:259±269 (1995).
47 Caseley JC and Coupland D, Environmental and plant factors
affecting glyphosate uptake, movement and activity, in The
Herbicide Glyphosate, ed by Grossbard E and Atkinson D,
Butterworths, London. pp 92±123 (1985).
48 Adkins SW, Tanipat S, Swarbrick JT and Boersma M, In¯uence
of environmental factors on glyphosate ef®cacy when applied
to Avena fatua or Urochloa panicoides. Weed Res 38:129±138
(1998).
49 McWhorter CG, Jordan TN and Wills GD, Translocation of 14C
glyphosate in soybeans (Glycine max) and johnson grass
(Sorghum halepense). Weed Sci 28:113±118 (1980).
50 Wills GD, Factors affecting the toxicity and translocation of
glyphosate in cotton (Gossypium hirsutum). Weed Sci 26:509±
513 (1978).
51 Marshall G, Herbicide-tolerant crops ± real farmer opportunity
or potential environmental problem? Pestic Sci 52:394±402
(1998).
52 O'Keefe MG and Makepeace RJ, Ef®cacy of glyphosate in arable
situations, in The Herbicide Glyphosate, ed by Grossbard E and
Atkinson D, Butterworths, London. pp 418±434 (1985).
53 Moreby SJ, The effects of herbicide use within cereal headlands
on the availability of food for arable birds. Proc Brighton Crop
Prot Conf±Weeds, BCPC, Farnham, Surrey, UK. pp 1197±
1202 (1997).
Pest Manag Sci 56:299±308 (2000)