Crop Protection 96 (2017) 30e43

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Crop Protection
journal homepage: www.elsevier.com/locate/cropro

A review of economic thresholds for invertebrate pests in UK arable

crops
M.W. Ramsden a, *, S.L. Kendall b, S.A. Ellis c, P.M. Berry c
a
RSK ADAS Ltd, Battle Gate Road, Cambridge CB234NN, United Kingdom
b
RSK ADAS Ltd, Gleadthorpe, Meden Vale, Manseld, Nottingham NG20 9PD, United Kingdom
c
RSK ADAS Ltd, High Mowthorpe Duggleby, Malton, North Yorkshire YO17 8BP, United Kingdom

a r t i c l e i n f o a b s t r a c t

Article history: The economic threshold for an invertebrate pest of an arable crop is the population density at which
Received 29 August 2016 control measures should be implemented to prevent economic damage. It is a valuable method of
Received in revised form determining whether or not control measures are necessary against an individual pest or group of pests.
16 January 2017
For thresholds to be effective, farmers and agronomists need to be condent that they accurately reect
Accepted 17 January 2017
the risk of economic loss in relation to the current cropping systems, as well as being practical to use. A
lack of condence can lead to the use of insurance sprays (insecticides applied irrespective of actual pest
abundance), which are environmentally damaging, increase the risk of pest resistance, and decrease
Keywords:
Crop tolerance
gross margins. We found that most current economic thresholds for pests of arable crops are not based
Invertebrate pests on published evidence, and almost none account for the ability of crops to tolerate pest damage, or the
Threshold amount, or type of crop damage that pests can cause. Furthermore, many of the methods of pest
Arable assessment are impractical, do not guarantee sufciently accurate estimates of pest abundance, and are
Pest damage not described with sufcient detail to ensure consistency of pest assessment. Following a critique of
current economic thresholds, this paper describes the relationship between invertebrate pest damage to
crops, yield formation, and crop tolerance to pest damage, and describes what crop information is
required to account for the capacity of crops to tolerate damage. This understanding is used to identify
the key elements of economic threshold schemes for arable invertebrate pests, and describe a process by
which thresholds can be applied. Finally we discuss the impact revised thresholds would have on crop
production, and the further work needed to develop accurate, reliable, practical economic threshold
schemes within integrated pest management strategies. We conclude that effective management of
invertebrate pests in arable crops is reliant on: (1) Quantifying the crop damage a pest can cause; (2)
Understanding the degree of tolerance a crop has to pest damage, and (3) Accurate and practical methods
for assessing pest abundance.
2017 Elsevier Ltd. All rights reserved.

Contents

1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
2. Current levels of adoption of economic thresholds in arable crops . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
3. Accounting for crop resistance and tolerance to pest damage . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
3.1. Yield formation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
3.2. Host-plant resistance to invertebrate pests . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
3.3. Passive and reactive tolerance to pests . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
4. Quantifying pest damage . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
5. Practicalities of assessing pest abundance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
6. Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
7. Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41

* Corresponding author.
E-mail address: mark.ramsden@adas.co.uk (M.W. Ramsden).

http://dx.doi.org/10.1016/j.cropro.2017.01.009
0261-2194/ 2017 Elsevier Ltd. All rights reserved.
 M.W. Ramsden et al. / Crop Protection 96 (2017) 30e43 31

Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41

1. Introduction are used to dene key aspects of threshold applications, developed

from denitions in Painter (1951), Stern et al. (1959) and Pedigo
Invertebrate pests can cause substantial yield losses in arable et al. (1986);
crops (Oerke, 2006; Culliney, 2014). Reductions in pesticide avail-
ability is of concern, because current pest management relies on
Crop damage e Crop injury which leads to measurable loss of
these products to protect crops from extensive damage (Hillocks, yield or reduction in quality.
2012). Insecticides are an important component in the manage-
Crop injury - The effect of pest feeding, or other activities, on the
ment of invertebrate pests, improving crop production when growth or appearance of crop plants.
applied correctly (Oerke and Dehne, 2004). The low cost of
Economic damage e The extent of crop damage that justies the
frequently used insecticides (e.g. synthetic pyrethroids) relative to cost of control.
potential economic losses associated with invertebrate pests en-
Economic injury level (EIL) e The lowest pest population density
courages prophylactic applications (e.g. calendar or insurance that will cause economic damage.
sprays) even though these are not always the most economic op-
Economic thresholde The pest population density at which
tion (Johnson et al., 2009; De Freitas Bueno et al., 2011; Reisig et al., control measures should be implemented to prevent pest pop-
2012). Prophylactic insecticide application can also have long term ulations reaching the EIL.
impacts, resulting in economic losses, compromised pest man-
Host-plant resistance e The relative amount of heritable quali-
agement and increased risk of damage to the surrounding envi- ties possessed by the plant which inuence the ultimate degree
ronment (Pimentel, 2005). of damage done by invertebrate pests.
Resistance is a constant threat to pesticide efcacy, and inap-
Host-plant tolerance e The capacity for a crop to be injured
propriate or excessive use increases the likelihood of resistance without any discernible impact on yield.
developing (Bass et al., 2014). Resistance to pyrethroids for example
has already been identied in several important pests, including Thresholds exist for most pests in the UK, however limited data
pollen beetle Meligethes aeneus F. (Coleoptera: Nitidulidae, Slater have been collected on their implementation, and few account for
et al., 2011), cabbage stem ea beetle Psylliodes chrysocephala L. variability in crop value or input costs over time. The Food and
(Coleoptera: Chrysomelidae, Hojland et al., 2015), grain aphids Environment Research Agency (FERA) oilseed rape (Brassica napus
Sitobion avenae F. (Hemiptera: Aphididae, Foster et al., 2014), and L.) survey and Insecticide Usage Survey show that, between 2004
the peach potato aphid Myzus persicae (Sulzer) (Hemiptera: Aphi- and 2014, the amount of insecticide applications on oilseed rape
didae, Bass et al., 2014). Resistance drives the market for new steadily increased, reecting both an increase in total weight of
chemicals, the development and registration of which cost sub- active ingredients applied, and an increase in average number of
stantial amounts over many years (Sparks, 2013). These costs are applications per year (PUS statistics 2004e2014). There is little
ultimately reected in pesticide prices to the farmer. Aside from evidence that farmers regularly use economic thresholds to sup-
these direct costs, inappropriate application of insecticides may port insecticide application decisions (Cohen et al., 1998), high-
reduce the abundance of benecial organisms such as naturally lighted for example, by the disparity between the proportion of
occurring predators of pests, leading to reductions in yield (Landis oilseed rape elds in which insecticides targeting pollen beetle
et al., 2000; Relyea, 2005; Desneux et al., 2007; Geiger et al., 2010; were applied and the proportion of those sites in which the eco-
Brittain and Potts, 2011). nomic threshold for the pest was exceeded (Fig. 1).
Concerns over the long term impacts of pesticide applications The availability of cheap pyrethroids, which offer a low cost and
have led to increased restrictions on use, removal of products from effective way of rapidly reducing pest numbers, is likely to
the market, and more rigorous procedures for registration of new encourage farmers to apply insecticides as an insurance spray and
active ingredients (Hillocks, 2012; Kohler and Triebskorn, 2013; van probably accounts for a signicant proportion of unnecessary ap-
der Sluijs et al., 2014). The European Commission has proposed plications (Pannell, 1991; Pedersen et al., 2012; Dewar, 2016).
more changes to the assessment of active ingredients, with the However, there are also a number of other reasons why insecticides
potential to reduce further the number of products available in are applied without reference to economic thresholds. Firstly, the
arable farming (Skevas et al., 2013). The EU Sustainable Use thresholds may be based on outdated or unsubstantiated research,
Directive (SUD), SUD 2009/128/EC, provides guidelines on the use and so are considered unreliable by farmers and agronomists (Ellis
of pesticides, promoting low input regimes, including Integrated et al., 2012). Secondly, thresholds may fail to account for variation
Pest Management (IPM) (Clarke et al., 2009). To comply with the in the crop's ability to resist and/or tolerate pest damage (Gu et al.,
SUD and to cope with the ongoing reductions in the availability of 2008). Thirdly, it may be too time consuming to undertake the
crop protection chemicals, farmers and agronomists need to give assessment methods required to determine if a threshold has been
much greater consideration to decisions concerning whether or not reached, making them impractical in comparison to the time and
to apply insecticides. This will require improved risk assessment, cost efciency of applying pesticides (Sharma et al., 2011; Ellis et al.,
fundamental to which is the use of robust, user friendly economic 2012). Given these concerns, and in the context of uctuations in
thresholds for invertebrate arable pests (Rose et al., 2016). the value of crops and the cost of plant protection products, farmers
Economic thresholds are a valuable method of assessing understandably err on the side of caution.
whether or not control measures, such as insecticides, are neces- Several authors have previously considered the main compo-
sary to reduce the risk of economic losses. They are usually dened nents that comprise a comprehensive economic threshold scheme
in terms of the number of a pest per unit area, per plant, or per part (Bardner and Fletcher, 1974; Poston et al., 1983; Pedigo et al., 1986;
of plant, above which action should be taken. The following terms Litsinger, 2009). In this review, we summarise current economic
32 M.W. Ramsden et al. / Crop Protection 96 (2017) 30e43
thresholds and highlight factors that may be responsible for the
poor uptake of thresholds at present. We then describe how yield is
formed in arable crops, and how this inuences the degree to which
crops can resist and tolerate crop injury. We outline the importance
of quantifying the damage caused by pests in terms of yield, and the
practicalities of pest assessment methods. Based on existing
studies, we consider the following as the key components for a
science-based economic threshold for arable invertebrate pests;
1) Economic thresholds should account for a crop's ability to resist
and/or tolerate pest injury, and the degree to which this varies
under normal conditions. This will identify crops not at risk of
yield loss, and those which require further assessment.
2) Economic thresholds should be based on known quantication
of a pest's potential to cause crop damage.
3) Economic thresholds must be based on one or more easily
assessed traits in the crop at a relevant growth stage.
4) Economic thresholds must account for the value of the crop and
of plant protection products, and reect how these change over
time.
We propose future research to help promote the adoption of
economic thresholds as a vital component of integrated pest
management, in order to minimise reliance on insecticides. We also
discuss strategies for improving pest assessment methods so that
they become more user friendly.
2. Current levels of adoption of economic thresholds in
arable crops
The invertebrate pests capable of causing signicant yield los-
ses, either through reductions in yield or quality, for the main
arable crops grown in the UK are summarised in Table 1. Eight of the
thirty four pests have no established threshold. The majority of the
thresholds identied have not been peer reviewed and most were
developed over twenty years ago, often based on anecdotal evi-
dence rather than robust scientic research. For example, the
threshold for treatment of infestations of cabbage seed weevils
Ceutorhynchus assimilis (Paykull) Coleoptera: Curculionidae in
oilseed rape is one weevil every other plant in northern UK, and one
weevil per plant elsewhere (Ellis et al., 2012). However, there is no
scientic or grey literature available describing how this gure was
calculated, raising questions over its accuracy and of other
Fig. 1. The proportion of sites monitored in which the population density of pollen
beetle exceeded the economic threshold, and the proportion of insecticide application
targeted at pollen beetle at these sites, in years surveyed between 2004 and 2014
(adapted from FERA pesticide usage survey).
unsubstantiated thresholds. Furthermore, there is no denition of
which regions constitute northern UK. Unpublished thresholds
rarely detail how assessments should be done, the number
required, or account for associated costs. Inconsistencies in
assessment methods increases the risk of misinterpretations,
leading to both the treatment of crops when the pest is not posing a
risk to yield, and failures to treat damaging pest infestations where
they do occur.
Eleven of the thresholds currently in use had been peer
reviewed, including that of the wheat bulb y, Delia coarctata
(Fallen) (Diptera: Anthomyiidae), an important pest of winter
wheat, Triticum aestivum L. The threshold for using treated seeds or
for applying insecticides to control this pest was described by
Gough et al. (1961). This was based on the knowledge of crop
physiology, ecology and the relative crop value and crop protection
costs of the time, and has not been reviewed since. Ongoing im-
provements in our understanding of factors such as the physio-
logical determination of yield enable better assessment of crop
tolerance to pests, yet updates to thresholds have been few and far
between. At best, current thresholds give a broad indication which
farmers can use to inform their pest management programme, at
worst they actively encourage prophylactic insecticide application.
Insecticide use is predicted to become increasingly restricted and
more expensive, and farmers are more likely to question whether
treatments should be applied (Sparks, 2013). Accurate economic
thresholds can help with this decision, but only where they accu-
rately reect risks to crop yield and account for current costs and
crop value.
3. Accounting for crop resistance and tolerance to pest
damage
The yield potential of a crop can be dened very simply as a
function of the light intercepted and radiation use efciency, the
product of which is biomass and the partitioning of biomass to yield
(Reynolds et al., 2009). Loomis and Amthor (1999) suggest that
achieving potential yield is conceptually simple e the extent and
duration of radiation interception must be maximised, the captured
energy must be used in efcient photosynthesis, new assimilates
must be partitioned in ways that provide optimal proportions of
leaf, stem, root and reproductive structures and these structures
must be maintained at minimum cost. Abiotic and biotic stresses
are partly responsible for the difference between potential yield
and actual yield, including damage by invertebrate pests. Soil
health plays an important underlying role in achieving yield po-
tentials, dictating the availability of water and nutrients to the crop
(Arshad and Martin, 2002; Kibblewhite et al., 2008). Factors such as
soil compaction can reduce resource availability, and this in turn
leads to reductions in yield (Hamza and Anderson, 2005) and af-
fects above ground interactions between the crop, its pests and
their natural enemies (Altieri and Nicholls, 2003; Garratt et al.,
2011; Butler et al., 2012).
3.1. Yield formation
For most crop plants, the relationship between crop injury and
yield loss varies with the growth stage of the plant at the time of
injury, the ability of the plant to resist and/or tolerate the injury,
and the part of the plant affected. This is normally related to
whether the yield of a plant is limited by the supply of photosyn-
thate (source limited), the capacity of the yield organ to accumulate
available carbohydrate (sink limitation) or a combination of both
(Shearman et al., 2005; Berry and Spink, 2006; Bingham et al.,
2007a, 2007b; Serrago et al., 2013). The yield of sink limited
crops is inuenced most by factors which impact before owering
 M.W. Ramsden et al. / Crop Protection 96 (2017) 30e43 33

Table 1
Current economic thresholds and methods of assessment for key UK arable pests.

Latin Common name Current Economic Threshold Threshold Assessment method Time per assessment (hrs)
peer
reviewed

Cereals
Agriotes spp. Wireworms Use seed treatment if > 750,000/ha. No Assess numbers in soil by 1.5 to sample, 1.5 to extract
Damage likely even with seed taking twenty 10 cm (depends on soil type
treatment if numbers >1.25 million/ha diameter x 15 cm deep
soil cores per 4 ha area
Set grain traps to attract 1.0 to set traps
wireworms 1.0 to examine
Chlorops pumilionis Gout y Eggs present on half of plants at No Examine plants for gout 0.5
GS12 for winter crops. No threshold y eggs
for spring crops
Delia coarctata Wheat bulb y >5 million eggs/ha e damage inevitable (1) Assess egg laying by taking 1.5 to sample, 1.5 to extract
2.5e5 million eggs/ha e damage likely 20, 10 cm diameter x 15 cm (depends on soil type)
<1.25 million eggs/ha e late sown deep soil cores/4ha
(NoveMar) crops may suffer damage
10% tillers attacked at GS20 Assess tiller infestation by 0.5 to sample, 1.0 to dissect
15% tillers attacked at GS21 dissecting 50 randomly
20% tillers attacked at GS23 chosen plants.
Deroceras reticulatum Grey eld slug Four or more slugs per refuge trap (2,3) Nine refuge traps or 13 for 0.5 to set traps, 0.5 to examine
elds larger than 20 ha in
a W pattern. Bait with
chicken layers mash
Haplodiplosis marginata Saddle gall midge NA NA NA NA
Opomyza orum Yellow cereal y >300 eggs/m2 (drilled before No No recognised assessment 0.5 to sample, 1.0 to dissect
mid-October) method. Can assess tiller
infestation as above
Oscinella frit Frit Fly >10% of plants showing No Can assess tiller infestation 0.5 to sample, 1.0 to dissect
Sitobion avenae, Summer aphids, 50% tillers infested before GS 61 66% (4,5) Visually examine 100 1
Metopolophium direct feeding damage; tillers infested from GS61 to two weeks randomly-chosen tillers
dirhodum Grain aphid & before end of grain lling for presence of aphids
Rose-grain aphid
Sitobion avenae, Autumn aphids, None - spray if aphids are present No Visually examine at least 1
Rhopalosiphum padi transmission of BYDV; 50 randomly chosen plants
Grain aphid & for aphids during
Bird cherry aphid September to November
Sitodiplosis mosellana Orange wheat >120 male midges/trap/day in (6,7,8,9) Assess soil for presence of 1.5 to sample,
blossom midge pheromone traps cocoons/larvae. Take
60 cores with 2.5 cm
cheese corer in W
pattern from area not
exceeding 15 ha
Feed crops e1 midge/3 ears Pheromone traps 0.5 to set, 0.5 to examine
Milling and seed crops e1 midge/6 ears Field monitoring by 0.5
parting crops and counting
midges disturbed.
Low risk <10 larvae and cocoons/kg soil Yellow sticky traps 0.5 to set, 0.5 to examine
Moderate risk 11e40 larvae and
cocoons/kg soil
High risk >41 larvae and cocoons/kg soil
Tipula paludosa, Leatherjackets >50 leatherjackets/m2 for spring cereals No Extraction in brine 1.5
Tipula oleracea solution. Drive 10 cm
diameter drainpipe
into soil and
part ll with brine.
Leatherjackets oat
to surface (19)
5 leatherjackets/m row for spring cereals Extraction using 0.5 to sample, 1.0 to extract
Blasdale apparatus (20)
Oilseed Rape
Brevicoryne brassicae Cabbage aphid Winter rape - >13% plants infested (10) Visually examine at least 0.5
before petal fall. Spring rape - >4% 50 randomly chosen plants
plants infested before petal fall for aphids in spring/early
summer
Ceutorhynchus assimilis Cabbage seed weevil >0.5 weevils/plant in northern Britain No Beat 25 randomly chosen 0.5
1 weevil/plant elsewhere plants over a white tray
Ceutorhynchus Cabbage stem weevil Treat if 2 weevils/plant of any species No Visual inspection of at 2 weevils/plant of any species
pallidactylus exceeded least 50 randomly
chosen plants for
any weevils
Deroceras reticulatum Grey eld slug One or more slugs per refuge trap (11,12) Nine refuge traps or 0.5 to set traps, 0.5 to examine
13 for elds larger than
20 ha in a W pattern.
Bait with chicken
layers mash
(continued on next page)
34 M.W. Ramsden et al. / Crop Protection 96 (2017) 30e43

Table 1 (continued )

Latin Common name Current Economic Threshold Threshold Assessment method Time per assessment (hrs)
peer
reviewed

Dasineura brassicae Brassica pod midge No

Meligethes aeneus Pollen beetle Less than 30 plants/m2 25 beetles No Beat 25 randomly 0.5
per plant 30e50 plants/m2 18 beetles chosen plants over
per plant 50e70 plants/m2 11 beetles a white tray
per plant More than 70 plants/m2 7
beetles per plant
Myzus persicae Peach potato None. Treat if aphids present No Visually examine at 1
aphid (TYV vectors) least 50 randomly
chosen plants for
aphids in autumn
Phyllotreta nemorum Large striped ea >25% leaf area eaten at 1e2 true No All ea beetles: 0.5
beetle leaf stage. Assess plants for
Phyllotreta spp. Turnip ea beetle >50% leaf area eaten at 3e4 true No presence of shot
leaf stage. holes
Psylliodes luteola Wessex ea beetle No
Psylloides Cabbage stem ea >35 beetles/water trap (13,14) All ea beetles: Yellow 0.5 to set, 0.5 to examine
chrysocephala beetle water traps (two in
headland, two in eld)
2 larvae/plant All ea beetles: Dissect 0.5 to sample, 1.5 to extract
up to 50 randomly
chosen plants for larvae
Beans
Aphis fabae Black bean aphid 10% plants infested at early owering, (15) Visually examine at 0.5
or 5% to prevent virus transmission least 50e100 randomly
chosen plants for aphids
on the south west
headland in spring/early
summer
Bruchus rumanus Bruchid beetles Spray when adults present and beans (16) Examine crops for 0.5
or bean beetle have rst pods on lowest trusses adults during owering
but only after temperature has by tapping the stems
reached 20C on two consecutive days. into your hand or a
Repeat in 7e10 days at receptacle
Ditylenchus dipsaci Stem nematode Zero tolerance for seed crops. No Soil cores 0.5
Seed checked for presence of
nematode. If found advice is
not to use it
Sitona lineatus Pea and bean weevil Treatment advised at rst signs No Visually examine at 0.5
of notching where there is a least 50 randomly
history of damage. chosen plants for
Cone trap available from Agralan weevils/leaf notching
Ltd with pheromone bait. Traps in spring/early summer
to be inspected every other day.
Threshold is an average of 30
adults per trap on a single occasion.
Peas
Acyrthosiphon pisum Pea aphid 15-20% plants infested at early ower. No Inspect about 50e100 0.5
Yield benets will be found up to plants between owering
4 podded trusses. and four trusses set for
aphids
Contarinia pisi Pea midge Susceptible crops are those which (17) In areas where midge 0.5e1.0
have reached enclosed bud stage. was a problem the
If adult midges can be found previous year, crops
an insecticide should be applied should be inspected
as soon as possible to reduce the when the oldest bud is
risk of eggs being laid. A pheromone about 6 mm long. Gently
system is available in the UK from squeeze leading shoots
Oecos Ltd. to immobilize midges.
Open buds and assess
for midges
Cydia nigricana Pea moth Combining peas - 10 or more moths (18) Set two pheromone 0.5/day
in any one of a pair of pheromone traps per eld and
traps on two consecutive occasions. count moths caught
Vining peas e spray at full ower if daily
any moths caught
Sitona lineatus Pea and bean weevil Treatment at rst signs of notching No As with beans above 0.5
where there is a history of damage.
Also cone trap, see above
Potatoes
Agriotes spp Wireworms None. Soil sampling will only detect No Assess numbers by 1.5 to sample, 1.5 to extract
down to 60,000/ha and damage is taking 20, 10 cm (depends on soil type)
possible below this level. diameter x 15 cm
deep soil cores/4ha
 M.W. Ramsden et al. / Crop Protection 96 (2017) 30e43
Table 1 (continued )
Latin Common name Current Economic Threshold
White PCN Globodera Potato cyst nematode Greater than 10 eggs/g soil. At egg
pallida, yellow populations greater than 60 eggs/g
PCN G rostochiensis soil growing potatoes is not
recommended.
Aphidoidea spp. Aphids None
Gastropoda spp. Slugs 1 slug/refuge trap over three day
period in late JulyeAugust
Linseed
Aphona euphorbiae Large ax ea beetle None
Longitarsus parvulus Small ax ea beetle None
Thrips angusticeps Field thrips None
(1) Gough et al., 1961; (2) Glen, 2005a; (3) Glen et al., 2006; (4) George and Gair, 1979; (5) Oakley and Walters, 1994; (6) Pivnick and Labbe
1989; (8) Larsson 1992; (9) Ellis et al., 2009; (10) Ellis et al., 1999; (11) Glen, 2005b; (12) Glen et al., 2006; (13) Walters et al., 2001; (14) Green, 2007, (15) Way et al., 1977; (16)
Ward and Smart, 2011; (17) Hilbur, 2001; (18) Wall, 1988. (19) Stewart and Kozicki, 1987; (20) Blasdale, 1974.
(when the sink size is determined), and source limited crops are
inuenced most by factors which impact after owering (when the
source size is determined). The yield of crops which are co-limited
by sink and source is affected by factors impacting on both pre- and
post-owering phases of growth. This means that there is potential
for the same pest to affect the yield of a sink or source limited crop
in different ways.
In sink limited crops, such as oilseed rape and barley, Hordeum
vulgare L., factors which restrict the determination of grains/m2 are
of great importance (Mendham et al., 1981; Berry and Spink, 2006;
Bingham et al., 2007ab, Fig. 2a). For example, in barley insufcient
shoot production has a large effect on potential yield (Bingham
et al., 2007ab). Pests which affect grain lling in these crops, such
as those that reduce canopy size after owering, are expected to
have a smaller effect on yield. Barley is particularly sensitive to
reductions in ear number because it has limited capacity to
compensate by increasing the number of grains per ear. In contrast
to barley, wheat can compensate for low ear numbers by increasing
grains per ear. Thus, whilst wheat can be co-limited by an insuf-
cient source and sink, wheat is generally considered as being source
limited in the UK environment (Shearman et al., 2005). As a result,
wheat has greater tolerance to reductions in shoot number, but a
lower tolerance to pests which reduce the supply of assimilate to
growing grains through reductions in green leaf area (Fig. 2b).
However, this could differ in other environments (Shearman et al.,
2005).
There will be exceptions to these general principles. Firstly,
environmental conditions or crop management may change a crop
from being sink to source limited, or vice-versa (Serrago et al.,
2013). A barley crop which produces an unusually large number
of ears, and as a result many grains/m2, may become more source
limited than sink limited (i.e. there is not enough source to ll all
the grains) (Bingham et al., 2007b). Secondly, the severity of the
impacting factor is important; a severe reduction in canopy size
during seed lling will reduce the yield of a sink limited crop
(Bingham et al., 2007b). Nevertheless, understanding whether a
crop is sink or source limited will generally be a useful indicator of
those phases of crop growth at which pest damage will have the
greatest effect on yield.
An important management tool for maximising crop yield is
optimisation of plant density, as this has a signicant impact on the
ability of individual plants to capture resources (Arduini et al.,
2006; Ellis and Berry, 2012). The relationship between plants/m2
35
Threshold Assessment method Time per assessment (hrs)
peer
reviewed
No Take soil sample comprising 0.5 to take sample
about 50 cheese corer cores
(approx 1e2 kg soil) and
submit to laboratory to
assess egg numbers/g soil
and cysts/100g soil
No Visually examine 50e100 0.5
plants for aphids
No Refuge traps baited with 0.5 to set traps, 0.5 to examine
chicken layers mash
No Visually examine 50e100 0.5
plants for beetles/damage
No Visually examine 50e100 0.5
plants for beetles/damage
No Visually examine 50e100 0.5
plants for thrips/damage
, 1993; (7) Kurppa and Husberg,
and yield can be explained in terms of the crop's ability to
compensate for low plant numbers by increasing shoot production,
grains per ear and individual grain weight. Yield is typically
increased by increasing seed rate or plant population up to a point,
and then reduced by super-optimal plant populations due to lod-
ging. Optimum crop density is affected by latitude, rotational po-
sition, sowing date, nitrogen nutrition, and weather (Spink et al.,
2000). Most crops are capable of achieving high yields from a
range of densities because individual plants adapt to local envi-
ronments. In wheat, optimum plant population can range from 70
plants/m2 to >600 plants/m2 (Whaley et al., 2000; Spink et al.,
2000). In oilseed rape, optimum plant population can range from
<10 plants/m2 to >160 plants/m2 (Roques and Berry, 2016).
A key physiological trait relating to sink and source limitation is
the ability of crops to maximise green leaf area. This directly relates
to the amount of light captured for photosynthesis, and ultimately
underpins yield formation. In oilseed rape, pod and seed survival
are related to the amount of radiation intercepted by photosyn-
thetic tissue per ower and per pod respectively (Leterme, 1988;
Mendham et al., 1981). In both wheat and barley, the potential
size of the grain is determined between ear emergence and early
grain development (Cochrane and Duffus, 1983; Bingham et al.,
2007b), and greater radiation and lower temperatures between
ear emergence and the start of grain lling are positively related to
nal grain weight (Bingham et al., 2007b). Similar principles apply
for other crops, including potatoes, Solanum tuberosum (L.), eld
beans, Vicia faba (L.), and sugar beet, Beta vulgaris (L.) (Weightman
et al., 2005; Allen et al., 2004; Jaggard et al., 2009).
Green Area Index, GAI is the ratio of green leaf area to the area of
ground in which the crop is growing. This is a fundamental factor in
yield formation, and as such it must be considered when assessing
the impact of pests. Plants have evolved various heritable mecha-
nisms to protect green leaf area and other tissues against injury
from invertebrate pests, broadly grouped into host-plant resistance,
and tolerance (Stout, 2013). Host-plant resistance, through antibi-
osis and antixenosis, and tolerance have become extremely
important aspects of crop variety breeding, making substantial
impacts in crop production over the last 60 years (Smith and
Clement, 2012; Stout, 2013). While both strategies fundamentally
serve a similar function, they differ in the mechanisms underlying
the response to invertebrate pest pressure and can provide com-
plementary protection against multiple pest attacks (Muola et al.,
2010).
Fig. 2. (a): Yield formation in UK barley is usually sink limited, as ears have limited capacity to compensate for low shoot numbers by increasing grain number. Excess production of
photo-assimilate cannot form additional yield, and increased yield is achieved primarily by increasing the number of ears per m2. Pest damage which reduces shoots/m2 in barley
will have a greater impact on yield than pest damage to green leaf area during grain lling.(b): Yield formation in UK wheat is usually source limited, as ears can have higher and
more exible grain numbers. A lack of green leaf area can fail to produce sufcient photo-assimilate to ll all potential grain sites. Increased yield is often achieved by maximising
green leaf area, which can fall short of the maximum grain capacity. Pest damage which reduces green leaf area, or which reduces assimilate transfer to the grains in wheat will have
a greater impact on yield than pests damage which reduces shoot number, providing shoot numbers remains above the threshold for yield potential (Table 2), because the crop
compensates by increasing grains per ear.
 M.W. Ramsden et al. / Crop Protection 96 (2017) 30e43
3.2. Host-plant resistance to invertebrate pests
Host-plant resistance reduces the colonisation or acceptance of
the crop by a pest (antixenosis; Painter, 1951; Kogan and Ortman,
1978), or reduces the growth, survival or fecundity of a pest
feeding on it (antibiosis) (Painter, 1951). Antixenosis is achieved
through mediated release of attractant and deterrent volatile
organic chemicals (VOCs) that manipulate the behaviour of pests
and their natural enemies (Palaniswamy and Lamb, 1993; Ferry
et al., 2004; Le Roux et al., 2010), and the creation of physical bar-
riers to pest attachment, feeding and oviposition, such as tri-
chomes, spinescence, waxy cuticles and sclerophylly (Mitchell
et al., 2016). Antibiosis can involve primary metabolites as well as
the production of secondary metabolites that reduce pest fecundity
or survival, often produced in response to damage (Bennett and
Wallsgrove, 1994; Hesler, 2005; Massey et al., 2006; Keeping and
Kvedaras, 2008; Bolton, 2009; Mitchell et al., 2016). In many in-
stances, antibiosis and antixenosis occur simultaneously; for
example, wheat cultivar resistance to orange wheat blossom midge,
Sitodiplosis mosellana (Ge hin, Diptera: Cecidomyiidae) is linked to
both reduced oviposistion on the wheat spikes (antixenosis; Lamb
et al., 2002) and production of phenolic acids which kill larvae
feeding in the grain (antibiosis; Ding et al., 2000).
The production of VOCs, physical barriers and secondary me-
tabolites requires additional resources. Therefore, crops experi-
encing resource stress exhibit lower resistance to herbivory than
crops that are not resource limited (Hanley et al., 2007; Mody et al.,
2008). Resistance may also vary over the course of the growing
season, depending on previous pest pressure, resource availability
and growth stage (Mody et al., 2008).
3.3. Passive and reactive tolerance to pests
Host-plant tolerance is the capacity for a plant to withstand or
recover from injury without any discernible impact on yield (Stout,
2013; Kock et al., 2016). Potato plants can generally tolerate around
20% leaf reduction at any life cycle stage with no discernible impact
on nal yield, and during some growth periods substantially more
leaf loss can be tolerated (Stieha and Poveda, 2015). There are ve
underlying physiological mechanisms for host-plant tolerance to
herbivory, broadly grouped as passive tolerance: high relative
growth rates, increased branching or tillering after release of apical
dominance, pre-existing high levels of carbon storage in roots, and
active tolerance: increased net photosynthetic rate after injure,
and the ability to reallocate carbon after injury from roots to shoots
(Sandras and Wilson, 1998; Strauss and Agrawal, 1999). Passive
mechanisms are based on excess production of tissue that can be
lost with no impact on yield (Table 2). The tissue damaged or
removed by pests is replaced by tissue that would otherwise have
been lost anyway through shedding or senescence (Sandras and
Wilson, 1998). In the case of potatoes, greater leaf area is pro-
duced than is needed to intercept all light (i.e. the crop is not source
limited), however, the quantity of excess leaf area changes over
time (Stieha and Poveda, 2015). Passive tolerance is dependent on
eld conditions such as crop density, crop variety and climatic
conditions, which inuence the quantity and quality of excess tis-
sue produced (Kays et al., 1974; Richards, 1988; Whaley et al., 2000;
Sparkes et al., 2006). In oilseed rape, the excess number of ower
buds produced is proportional to plant density, and so subsequent
tolerance of bud loss to pollen beetles M. aeneus also varies with
crop density (Trumble et al., 1993; Ellis and Berry, 2012). Many
other crops also show plant density dependent tissue production
which would impact the plant's capacity for passive tolerance
(Nienhuis and Singh, 1984; Reese et al., 1994; Diepenbrock, 2000;
Oliveira, 2000; Lopez-Bellido et al., 2005).
37
Passive tolerance only protects plants against pest attacks where
excess tissue is available to be lost. Where passive tolerance is not
possible, active mechanisms allow plants to reach their yield po-
tential despite herbivory (Reese et al., 1994; Sadras, 1995). Active
tolerance does not affect the behaviour, reproduction, growth, or
development of the attacking pests (Reese et al., 1994), but occurs
where plants replace or repair damaged tissue or redistribute re-
sources within the plant to compensate for the damage with no net
reduction in yield potential (Sadras, 1995). These mechanisms are
less well understood than passive actions, though some examples
are well described (Rubia et al., 1996). In winter wheat, gout y,
Chlorops pumilionis (Bjerkander) (Diptera: Chloropidae), attacks are
not considered to be of great importance as larvae feed only on a
single tiller prior to stem extension. The attacked plant can actively
divert resources into remaining tillers, and the nal yield is not
affected (Oakley et al., 1990). In spring sown wheat, however, gout
y attacks can have a greater impact as the wheat has less time to
produce additional tillers or to divert resources away from infested
stems prior to stem extension (Oakley et al., 1990). Under certain
conditions, active tolerance can lead to overcompensation where
pest action stimulates growth and/or resource redistribution that
increases yield, though this is uncommon in arable crops (Bardner
and Fletcher, 1974; Litsinger, 2009; Lebon et al., 2014). Active
tolerance is only functional where resource redistribution does not
divert resources away from yield formation, i.e. in crops that are not
sink limited.
As passive and active tolerance mechanisms will vary with
changing plant physiology, the impact that a pest has on yield
changes at different crop growth stages, different life stages of the
pest, under different eld conditions, and between crop types and
varieties. For example, infestations of adult cabbage stem ea
beetle, Psylliodes chrysocephala L. (Coleoptera: Chrysomelidae), can
lead to total crop failure in oilseed rape during crop emergence.
Their larvae can also cause some crop damage in very high numbers
during vegetative growth, but infestations of newly emerged adults
the following spring will inict practically no economic damage
during reproductive growth (Lamb, 1989; Hojland et al., 2015).
4. Quantifying pest damage
Understanding how different pests cause plant injury is
important for understanding how they may impact on crop yield
and quality. Several types of pest injury to plants have been pro-
posed by Boote et al. (1983), and these are adapted below for UK
arable pests.
1. Tissue consumers; e.g. slugs, ea beetles, pea and bean weevil,
pollen beetle, seed weevil.
2. Leaf senescence accelerators; e.g. aphids.
3. Stand reducers; e.g. slugs, wireworms, leatherjackets, dipterous
stem borers.
4. Photosynthetic rate reducers; e.g. aphid honeydew and sooty
moulds, leaf miners, leaf beetles.
5. Assimilate sappers; e.g. aphids, saddle gall midge, orange wheat
blossom midge.
6. Turgor reducers; e.g. nematodes, leatherjackets, wireworms
through root damage.
The rst four injury types affect light capture, while the
remaining two affect hydration and resource movement. A single
invertebrate pest may inict more than one type of injury. Tissue
removers usually do more damage than assimilate sappers, because
the plant must allocate resources for tissue replacement as well as
photosynthate (Litsinger, 2009).
The amount of injury an individual pest causes a plant, and how
38 M.W. Ramsden et al. / Crop Protection 96 (2017) 30e43

Table 2
Physiological potential of UK arable crops to tolerate pest damage.

Crop parameter Minimum to achieve Range in practice Typical degree of Source

potential yield tolerance to damage

Winter wheat
Plants/m2 <100 early sown 50 to 600 High Whaley et al. (2000)
200 late sown
2 2
Ears/m 400 400 to >2000 shoots/m High Spink et al. (2000)
GAIa at owering 5 to 7 5 to 10 Moderate AHDB Wheat Growth Guide
Post-owering Unknown Unknown Low
photo-assimilate
Winter barley
Plants/m2 Unknown Up to 600 Low/Moderate AHDB Barley Growth Guide
Ears/m2 Unknown 400 to >2000 shoots/m2 Low Bingham et al. (2007a,b)
GAI at owering 5 to 7 4 to 9 Low AHDB Barley Growth Guide
Post-owering Unknown Unknown Moderate
photo-assimilate
Spring barley
Plants/m2 Unknown Up to 600 Low AHDB Barley Growth Guide
Ears/m2 Unknown Unknown Low
GAI at owering Unknown Unknown Low
Post-owering Unknown Unknown Moderate
photo-assimilate
Winter oilseed rape
Plants/m2 Unknown 20 to 120 Moderate Roques and Berry (2016)
Pods/m2 7000e8000 5000e12,000 Moderate Berry and Spink (2006)
GAI at owering 3 to 4 3 to 6 Moderate Berry and Spink (2006)
Post-owering Unknown Unknown Moderate
photo-assimilate
Spring oilseed rape
Plants/m2 Unknown 20 to 120 Moderate Ellis and Berry (2012)
Pods/m2 7000e8000 Unknown Low Ellis and Berry (2012)
GAI at owering 3 to 4 Unknown Low Berry and Spink (2006)
Post-owering Unknown Unknown Moderate
photo-assimilate
Potatoes
Plants/m2 Market dependent Market dependent Low
GAI 4 3 to 8 Moderate Allen et al. (2004)
Peas
Plants/m2 Unknown Unknown Unknown
GAI 4 3 to 9 Low Weightman et al. (2005)
Beans
Plants/m2 10 to 100 10 to 100 Moderate Weightman et al. (2005)
GAI 4 3 to 9 Moderate/High Weightman et al. (2005)
a
GAI e green area index.

this in turn impacts yield formation is fundamentally important in
calculating economic thresholds. For a given pest, economic
thresholds can be calculated from; the injury attributable to an
individual pest, the amount of subsequent damage the plant can
tolerate without impact on yield (tolerance), the number of pests
per plant, the value of the crop, and the cost of applying treatments.
Pest damage can have a direct or indirect impact on crop yield,
depending on the crop character being attacked (e.g. leaf area,
shoots per m2, marketable yield per m2, Table 3). Where yield
forming (sink) tissue is attacked, quantifying subsequent yield
losses can be relatively straight forward. In the case of the pea moth
Cydia nigricana F. (Lepidopptera: Tortricidae), the larvae feed
directly on peas inside the pod. Yield loss per larvae can be esti-
mated and related to adult abundance, monitored using phero-
mone traps (Huusela-Veistola and Jauhiainen, 2006). Where pest
damage has an indirect impact on yield (i.e. on source or supportive
tissue), losses are more difcult to quantify and are often only
possible at a population level rather than per insect (Godfrey and
Wood, 1998; Larsson, 2005; Beckendorf et al., 2008). More
research is required to quantify the indirect damage associated
with loss of green leaf area as a result of pest damage.
Economic thresholds are based on a number of assumptions,
such as low variation in the market value of crop and the cost of
insecticide treatment over time. When these change dramatically
(e.g. if insecticides were to become ten times more expensive),
existing economic threshold may no longer be valid. This
emphasizes the need for peer review in dening economic
thresholds, so that all assumptions are published and better guid-
ance on their use is available to growers. It also highlights the need
for periodic reviews of established economic thresholds to ensure
they are still relevant to present market values.
5. Practicalities of assessing pest abundance
Economic thresholds need to be reliable, practical, and cost
effective. They can be difcult to calculate, often requiring detailed
knowledge of how an individual pest causes damage to a plant, the
impact that damage has on the nal yield, and the associated cost.
Economic thresholds for a pest often need to be transformed into a
more easily measurable variable in the eld, such as a quantica-
tion of damage or of pest abundance at an earlier life stage.
Over estimation of pest abundance and subsequent unnecessary
applications of insecticide can have long term detrimental effects
on yield through selection of resistant pests and damage to bene-
cial organisms. Where pest abundance is under estimated this can
lead to signicant losses in yield that could have been prevented.
The latter has a much more immediate impact on farming prots,
and drives the use of insurance sprays and limits use of thresholds,
the former is a growing threat to crop production in the UK. The
contribution of peer reviewed scientic research is fundamental in
ensuring that thresholds are robust, however, greater engagement
is needed with industry to ensure they are also practical. At present,
 M.W. Ramsden et al. / Crop Protection 96 (2017) 30e43 39

Table 3
Summary of the crop character affected by key pests and limitations in current knowledge.

Pest Crop character affected Method for assessing Minimum crop character Is pest damage Damage caused
crop character? to achieve potential yield quantiable? by single pest

Wheat
Grey eld slug Leaf area or plants/m2 Yes Known Yes Unknown
BYDV vectors General growth No Unknown Difcult Unknown
Gout y Shoots/m2 No Known Yes Known (1 shoot)
Wheat bulb y Shoots/m2 No Known Yes Unknown
Yellow cereal y Shoots/m2 No Known Yes Known (1 shoot)
Wireworms Plants/m2 Yes Known Yes Unknown
Leatherjackets Plants/m2 Yes Known Yes Unknown
Orange wheat blossom midge Seed lling No Unknown Difcult Unknown
Summer aphids Seed lling No Unknown Difcult Unknown
Saddle gall midge Seed lling No Unknown Yes Unknown
Barley
Grey eld slug Plants/m2 Yes Unknown Yes Unknown
BYDV vectors General growth No Unknown Difcult Unknown
Gout y Shoots/m2 No Known Yes Known (1 shoot)
Wheat bulb y Shoots/m2 No Known Yes Unknown
Yellow cereal y Shoots/m2 No Known Yes Known (1 shoot)
Wireworms Plants/m2 Yes Known Yes Unknown
Leatherjackets Plants/m2 Yes Known Yes Unknown
Summer aphids Seed lling No Unknown Difcult Unknown
Saddle gall midge Seed lling No Unknown Yes Unknown
Oilseed
Grey eld slug Plants/m2 Yes Known Yes Unknown
Summer aphids General growth No Unknown Difcult Unknown
Cabbage stem ea beetle Plants/m2 Yes Known Yes Unknown
Wessex ea beetle Plants/m2 Yes Known Yes Unknown
Turnip ea beetle Plants/m2 Yes Known Yes Unknown
Large striped ea beetle Plants/m2 Yes Known Yes Unknown
Pollen beetle Pods/m2 No Known Yes Known (9 pods)
Cabbage stem weevil General growth No Unknown Difcult Unknown
Cabbage seed weevil Seeds/m2 N/A Known Yes Known
Brassica pod midge Seeds/m2 N/A Known Yes Known
Cabbage aphid Seed lling No Unknown Difcult Unknown
Beans
Pea and bean weevil Leaf area or plants/m2 No Unknown Yes Unknown
Black bean aphid General growth No Unknown Difcult Unknown
Stem nematode Seeds/m2 No Unknown Yes Unknown
Bruchid beetles or bean beetle Marketable seed/m2 No Unknown Yes Unknown
Peas
Pea and bean weevil Leaf area or plants/m2 No Unknown Yes Unknown
Pea aphid General growth No Unknown Difcult Unknown
Pea moth Marketable yield/m2 No Unknown Yes Unknown
Potatoes
Potato cyst nematode Tubers/m2 No Unknown Yes Unknown
Wireworms Marketable yield/m2 No Unknown Yes Unknown
Aphids General growth No Unknown Difcult Unknown
Slugs Marketable yield/m2 No Unknown Yes Unknown
Linseed
Large ax ea beetle Plants/m2 Yes Unknown Yes Unknown
Small ax ea beetle Plants/m2 Yes Unknown Yes Unknown
Field thrips Plants/m2 Yes Unknown Yes Unknown

some assessment methods involve the use of bulky technical
equipment, such as the Salt and Hollick extraction of wheat bulb y
eggs and wireworms (Salt and Hollick, 1944), or Blasdale extraction
of leatherjackets (Blasdale, 1974) which are not feasible for farmers
or agronomists to use. However, these techniques are effective at
estimating pest numbers, and are used as part of annual surveys to
create regional risk maps. These risk maps provide a regional
mechanism for alerting growers to potential threats, or to the
arrival of pests which can then be related to the growth stage in
individual elds.
Above all, thresholds must be demonstrably cost effective. Their
role in pest management is undermined where the time taken or
the cost of equipment make it uneconomic to undertake the fre-
quency of assessments required. The distribution of most pests is
patchy rather than uniform, and assessments need to be done in
multiple locations across a eld. Current sampling methods avail-
able in UK arable pest management are summarised in Table 1. On
farms with several crops in rotation, these can quickly amount to a
signicant undertaking, even when several assessment can be done
concurrently. Where threshold assessments are overly complicated,
or incur excessive costs, it is unlikely that they will adopted.
6. Discussion
The current economic thresholds outlined in Table 1 do not
adequately provide growers with reliable, usable guidance on when
to apply insecticide treatments in the UK. New economic thresh-
olds are under development, such as the recently revised economic
threshold for pollen beetle in oilseed rape (Ellis and Berry, 2012),
though many established thresholds remain unsubstantiated. The
lack of scientically robust thresholds, failure to account for vari-
ation in crop resistance and/or tolerance, and difculties in
assessing pest numbers reduce grower condence in thresholds as
part of IPM in arable crops. Alongside this, a lack of transparency on
40 M.W. Ramsden et al. / Crop Protection 96 (2017) 30e43
how economic thresholds account for uctuations in crop value
and input costs further reduces their appeal to farmers (Rose et al.,
2016). Widespread prophylactic insecticide use is not sustainable
and chemical control is likely to become more restricted and
expensive in the future. Farmers are willing to adopt new strategies
to improve crop protection that reduce reliance on insecticides, but
need assurance that effective pest control will be maintained
(Bailey et al., 2009; Geiger et al., 2010; Lechenet et al., 2014). Eco-
nomic thresholds provide a method by which unnecessary insec-
ticide applications can be avoided and enable farmers to compare
the impact of alternative treatments on their crops. However, they
must be updated periodically to account for advances in crop
physiology, understanding of crop-pest interactions, and
agronomy. Insecticides will continue to play a key role within
management programmes, but, in order to maximise the longevity
of products, the frequency of application will need to be reduced.
Economic thresholds provide a framework for achieving this.
We have identied three key factors underpinning the devel-
opment of robust invertebrate economic thresholds; the crop's
ability to resist and/or tolerate pest injury, quantication of the
subsequent amount of damage an individual pest causes, and the
practicality of assessing pest numbers. Host-plant resistance and/or
tolerance is a vital, yet mostly absent, component of economic
thresholds, and has the greatest potential for rationalising insecti-
cide use. By improving our understanding of how resistance and
tolerance relate to crop growth and resource availability, we will be
better able to predict situations in which pest induced economic
damage is likely. As it is usually far easier to assess crop charac-
teristics or resource availability than pest abundance, crop assess-
ments could be developed that provide a quick method for
determining whether or not control measures are required. For
example, Oilseed rape produces signicantly more owers than the
optimum pod number required for potential yield, so there is an
excess number of owers per plant which could be sacriced to
pollen beetle attack without yield loss, which is inversely propor-
tional to the plant population density (Ellis and Berry, 2012).
Therefore, by measuring plant numbers in late winter, it is possible
to determine the number of excess buds a crop will produce. As
each pollen beetle is thought to consume nine buds (Ellis and Berry,
2012), the number required to consume all excess buds can be
calculated. If the number of beetles required to consume all excess
buds is not reached in oilseed rape crops, it is very unlikely that
crop injury will exceed the EIL, making any treatments applied
uneconomical (Ellis and Berry, 2012). It is therefore possible to
make decisions on the need for control measures well in advance of
the pest arriving in the crop. This is particularly cost effective as
more time can be spent on other crops which are less tolerant of
pest damage.
In many crops, only certain growth stages are vulnerable to
damage, and this provides assurance to farmers that control is not
needed before or after these periods. Extension of this concept to
include crop growth stages which may be vulnerable, but which
can resist and/or tolerate a greater quantity of pest damage than is
likely to occur, would further support farmers in reducing inputs
where yield loss risks are lower. To achieve this, more information
is needed on how crop characters (e.g. plant population density,
seed lling duration, seed/fruit/tuber number per plant, or leaf
area) relate to nal yield for a wider range of crop species than is
currently known. Combined with the ability to quantify the abun-
dance of a pest, this provides the foundations for tolerance related
thresholds. However, much of the information needed to create
these thresholds is not currently known (Table 3).
Economic thresholds need to be accurate, reliable, and simple to
understand. They must also relate to easily quantiable factors that
can be assessed quickly and cost effectively. In addition, in order
that economic thresholds remain relevant to farmers re-
quirements, they must account for changes in crop value and cost of
treatments over time. Over simplication can reduce the credibility
of thresholds, but complicated or time consuming assessments are
not practical on farms covering large areas with multiple crops. The
development of remote sensing technology to estimate crop
growth and condition, combined with models predicting the arrival
of pests, may enable farmers to identify risks to crop yield across
landscapes more effectively (Jurdak et al., 2015). Such technology
will be essential in providing low cost assessments, however, these
will not be possible without a more detailed understanding of the
underlying physiology of crop yield formation, pest behaviour, and
their interaction.
Deciding not to apply an insecticide is one of the most difcult
decisions a farmer or agronomist can make. Subsequent pest at-
tacks resulting in nancial losses can severely damage an agrono-
mist's reputation. In contrast, there is little chance of repercussions
against decisions to treat when no pest threat was present or would
have arisen. There is general acknowledgement for the need to
reduce agricultural inputs while maintaining crop yields, and this is
a growing challenge. By their nature, economic thresholds are es-
timates rather than absolutes, and so always carry an element of
uncertainty. The assumption thresholds make is that damage
causing invertebrates will only become pests above a certain pop-
ulation size. Aside from the main limitations discussed in this pa-
per, a few additional factors will need to be addressed in the future.
Firstly, pest populations are rarely evenly distributed within a eld,
but are patchy and dynamic so the economic threshold could be
exceeded in parts of a crop but not others (Fievet et al., 2007;
Vialatte et al., 2007). To detect this would inevitably involve mul-
tiple sampling points across a eld, so there is a trade-off between
eld scale accuracy and the time required to undertake assess-
ments (Miller, 2003; Petrovskii et al., 2014). Secondly, thresholds
should be used as part of IPM programmes, which incorporate a
range of control measures alongside insecticide treatment (Zhang
and Swinton, 2009). Thirdly, little is known about how damage
by one pest changes crop resistance and/or tolerance to subsequent
attacks by the same or different pests (Dangles et al., 2009).
Based on the ndings of this review, we propose that effective
thresholds must account for the ability of crops to resist and/or
tolerate pest injury, should be based on quantitative data on the
level of subsequent damage caused by pests, must be user friendly,
and must come with sufcient guidance on how they have been
produced, including any economic or ecological assumptions made.
There are two essential stages for the use of economic thresholds in
pest management (Fig. 3). The rst stage is deciding whether a crop
is at a growth stage which is vulnerable to pests, and whether the
pests are likely to be present. This is most effectively done remotely
by combining knowledge of crop physiology and pest ecology, crop
development and pest emergence modelling, complimented by
using remotely sensed data and in-eld assessments where
possible (Leather et al., 1989; Jacquemin et al., 2014). Where it is
conrmed that the crop is at risk, the second stage is to establish
the potential for damage to exceed economic injury levels. This may
require additional information about the crop, such as plant pop-
ulation density, green leaf area, previous pest incidence, manage-
ment, crop value, and the projected cost of applying a treatment.
This information can be gathered relatively quickly, and new
technology is becoming available making this information acces-
sible through online assessments of crop photographs, or high
resolution remotely sensed images of the crop (e.g. satellite im-
ages). Economic thresholds and previous knowledge of likely pest
abundance may at this point be sufcient to support no further
action, or alternatively could require an additional ineld assess-
ments to conrm whether or not treatment is required.
 M.W. Ramsden et al. / Crop Protection 96 (2017) 30e43
Fig. 3. Decision ow for management of arable invertebrate pests.
7. Conclusion
Regrettably, the rst question often asked about any pest
infestation is, what can be used to control it? rather than, does it
need to be controlled? Agronomists are able to identify plant
injury, however there is a lack of peer reviewed evidence about
how plant injury relates to yield. All too frequently efforts
concentrate on control measures rather than investigating the
interaction between the pest and the crop. Given the relative low
cost of pesticides, it is understandable that when farmers and
agronomists are faced with a potential yield loss, initial consider-
ation is given to which chemical will give the best control. However,
this does little to advance our knowledge of pest management or
our understanding of whether the pest is likely to impact on yield.
This review suggests that a failure to account for the variable
resistance and tolerance of crops to pest injury and subsequent
damage undermines the ability of farmers and agronomists to
accurately predict the cost effectiveness of treatments. Economic
41
thresholds that do not account for this are increasingly being
abandoned, and we suggest that many need to be revised to
improve their accuracy and value to IPM strategies. In the future,
improvements in the understanding of crop physiology and agro-
nomic techniques will further inuence threshold calculations,
requiring that they be under constant review. Reliable, repeatable,
practical thresholds are vital for ensuring pesticides are applied
only where needed.
Acknowledgements
This work was supported by the Department for Environment,
Food and Rural Affairs, United Kingdom [grant PS2814]. We are
grateful to FERA for provide data on pesticide usage and pollen
beetle data in the UK, and Kate Storer and staff at AHDB for their
comments in preparing this review.
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