Soil & Tillage Research 72 (2003) 203–211

Soil fertility management and insect pests: harmonizing

soil and plant health in agroecosystems
Miguel A. Altieri∗ , Clara I. Nicholls
Department of Environmental Science, Policy and Management, Division of Insect Biology,
University of California at Berkeley, Berkeley, CA 94720-3112, USA
Received 5 March 2002; accepted 3 March 2003

Abstract
Cultural methods such as crop fertilization can affect susceptibility of plants to insect pests by altering plant tissue nutrient
levels. Research shows that the ability of a crop plant to resist or tolerate insect pests and diseases is tied to optimal physical,
chemical and mainly biological properties of soils. Soils with high organic matter and active soil biology generally exhibit
good soil fertility. Crops grown in such soils generally exhibit lower abundance of several insect herbivores, reductions that
may be attributed to a lower nitrogen content in organically farmed crops. On the other hand, farming practices, such as
excessive use of inorganic fertilizers, can cause nutrient imbalances and lower pest resistance. More studies comparing pest
populations on plants treated with synthetic versus organic fertilizers are needed. Understanding the underlying effects of why
organic fertilization appears to improve plant health may lead us to new and better integrated pest management and integrated
soil fertility management designs.
© 2003 Elsevier Science B.V. All rights reserved.
Keywords: Soil fertility; Crop nutrition; Pest attack; Insect populations; Pest management

1. Introduction insects and diseases are reduced with organic farming

(Merrill, 1983; Oelhaf, 1978). Although this view is
Many researchers have suggested that increasing in- widespread, there have been surprisingly few attempts
sect pest and disease pressure in agroecosystems is due to test its validity. The few conducted studies sug-
to changes that have occurred in agricultural practices gest that lower pest pressure in organic systems could
since World War II. For example, the usage of fertiliz- result from the greater use of crop rotation and/or
ers and pesticides has increased rapidly during this pe- preservation of beneficial insects in the absence of pes-
riod and evidence suggests that such excessive use of ticides (Lampkin, 1990). Alternatively, reduced sus-
agrochemicals in conjunction with expanding mono- ceptibility to pests may be a reflection of differences in
cultures has exacerbated pest problems (Conway and plant health, as mediated by soil fertility management
Pretty, 1991). On the other hand, proponents of alter- (Phelan et al., 1995). Many researchers and also prac-
native agricultural methods contend that crop losses to ticing farmers have observed that fertility practices
that replenish and maintain high soil organic matter
∗ Corresponding author. Tel.: +1-510-642-9802;
and that enhance the level and diversity of soil macro-
fax: +1-510-642-7428.
and microbiota provide an environment that through
E-mail addresses: agroeco3@nature.berkeley.edu (M.A. Altieri), various processes enhances plant health (McGuiness,
nicholls@uclink.berkeley.edu (C.I. Nicholls). 1993).

0167-1987/03/$ – see front matter © 2003 Elsevier Science B.V. All rights reserved.
doi:10.1016/S0167-1987(03)00089-8
204 M.A. Altieri, C.I. Nicholls / Soil & Tillage Research 72 (2003) 203–211

Fig. 1. The potential synergism between soil fertility management and IPM.

Despite the potential links between soil fertility and cific pest populations. Soil fertility practices can im-
crop protection, the evolution of integrated pest man- pact the physiological susceptibility of crop plants to
agement (IPM) and integrated soil fertility manage- insect pests by either affecting the resistance of indi-
ment (ISFM) have proceeded separately. The integrity vidual plant to attack or by altering plant acceptability
of the agroecosystem relies on synergies of plant diver- to certain herbivores. Some studies have also docu-
sity and the continuing function of the soil microbial mented how the shift from organic soil management
community, and its relationship with organic matter to chemical fertilizers has increased the potential of
(Altieri and Nicholls, 1990). Most pest management certain insects and diseases to cause economic losses.
methods used by farmers can be considered soil fer-
tility management strategies and vice versa. There are
positive interactions between soils and pests that once 2. The effects of fertilization on plant resistance
identified can provide guidelines for optimizing to- to insect pests
tal agroecosystem function (Fig. 1). Increasingly, new
research is showing that the ability of a crop plant Studies of plant resistance to insect pests have
to resist or tolerate insect pests and diseases is tied shown that resistance varies with the age or growth
to optimal physical, chemical and mainly biological stage of the plant (Slansky, 1990). This suggests that
properties of soils. Soils with high organic matter and resistance is linked directly to the physiology of the
active soil biology generally exhibit good soil fertility plant and thus any factor that affects the physiology
as well as complex food webs and beneficial organ- of the plant may lead to changes in resistance to insect
isms that prevent infection. On the other hand, farm- pests.
ing practices that cause nutrition imbalances can lower Fertilization has been shown to affect all three
pest resistance (Magdoff and van Es, 2000). categories of resistance proposed by Painter (1951):
Much of what we know today about the relationship preference, antibiosis, and tolerance. The obvious
between crop nutrition and pest incidence comes from morphological responses of crops to fertilizers, such
studies comparing the effects of organic agricultural as changes in growth rates, accelerated or delayed ma-
practices and modern conventional methods on spe- turity, size of plant parts, and thickness and hardness
 M.A. Altieri, C.I. Nicholls / Soil & Tillage Research 72 (2003) 203–211 205

of epicuticle, also influence the success of many pest of organic and synthetic fertilizer effects on the nu-
species in utilizing the host. For example, Adkisson tritional content of four vegetables reported that the
(1958) reported nearly three times as many boll weevil OG vegetables consistently contained lower levels
larvae (Anthonomus grandis) on cotton (Gossypium of nitrate and higher levels of potassium, phospho-
hirsutum) receiving heavy applications of fertilizers rus, and iron than CG vegetables. The above studies
compared to unfertilized checks. He attributed these suggest that the lower foliar content of NO3 -N of
differences to the prolonged growing season for cot- OG crops may be a key factor in determining lower
ton resulting from the fertilizer amendment by which insect damage on crops fertilized with organic amend-
plants remain succulent longer and fruit later in the ments.
season than normal. Klostermeyer (1950) reported
that nitrogen fertilizer increased husk extension and
tightness of husks on sweet corn (Zea mays) influenc- 3. Nitrogen effects
ing corn earworm (Heliothis zea) infestation levels.
Meyer (2000) argues that soil nutrient availability The indirect effects of fertilization practices acting
not only affects the amount of damage that plants re- through changes in the nutrient composition of the
ceive from herbivores but the ability of plants to re- crop have been reported to influence plant resistance
cover from herbivory; however, these two factors are to many insect pests. Among the nutritional factors
rarely considered together. Describing the effects of that influence the level of arthropod damage in a crop,
soil fertility on both the degree of defoliation and com- total nitrogen (N) has been considered critical for both
pensation for herbivory for Brassica nigra plants dam- plants and their consumers (Mattson, 1980; Scriber,
aged by Pieris rapae caterpillars, Meyer (2000) found 1984; Slansky and Rodriguez, 1987).
that the percentage defoliation was more than twice In most studies evaluating aphid and mite response
as great at low compared to high fertility, even though to N fertilization, increases in N rates dramatically
plants grown at high soil fertility lost a greater abso- increased aphid and mite numbers. According to van
lute amount of leaf area. At both low and high soil Emden (1966) increases in fecundity and develop-
fertility, total seed number and mean mass per seed of mental rates of the green peach aphid, Myzus persicae,
damaged plants were equivalent to those of undam- were highly correlated to increased levels of soluble
aged plants. Thus soil fertility did not influence plant N in leaf tissue. Several other authors have also in-
compensation in terms of maternal fitness. dicated increased aphid and mite populations from
Effects of soil fertility practices on pest resis- N fertilization (Tables 1 and 2). Herbivorous insect
tance can be mediated through changes in nutritional
content of crops. At equivalent amounts of applied Table 1
nitrogen (100 and 200 mg per pot), Barker (1975) Summary of effects of inorganic fertilizers on mite abundance
found that NO3 -N concentrations in spinach leaves from selected studies (Luna, 1988)
(Spinacia oleracea L.) were higher when receiv- Nutrients Mite species Crop Numerical
ing ammonium nitrate than in plants treated with response
five organic fertilizers. In a comparative study of of insecta
midwestern USA conventional and organic farmers, N Panonychus ulmi Apple +
Lockeretz et al. (1981) reported organically grown N Tetranychus telarius Apple +
(OG) corn to have lower levels of all amino acids N T. telarius Beans +
N, P, K Two-spotted spider mite Beans/peaches +
(except methionine) than conventionally grown (CG) N T. telarius Tomato −
corn. Eggert and Kahrmann (1984) also showed CG N, P T. telarius Apples +/−
dry beans (Phaseolus vulgaris) to have more pro- N, K Bryobia praetiosa Beans +/−
tein than OG beans. Consistently higher N levels in N, Ca Heliothrips Beans +/−
the petiole tissue were also found in the CG beans. haemorrhoidalis
a Symbols: (+) increase in density with increasing rates of
Potassium and phosphorus levels, however, were
fertilizer element; (−) decrease in density with increasing rates of
higher in the OG beans petioles than in the CG beans. fertilizer element. Slash separates the effects of fertilizer elements
Schuphan (1974) in a long-term comparative study listed in nutrients column.
206 M.A. Altieri, C.I. Nicholls / Soil & Tillage Research 72 (2003) 203–211

Table 2 sect populations found to increase following chemical

Summary of effects of inorganic fertilizers on aphid abundance N fertilization included fall armyworm in maize, corn
from selected studies (Luna, 1988)
earworm on cotton, pear psylla on pear (Pyrus sp.),
Nutrients Insect species Crop Numerical Comstock mealybug (Pseudococcus comstocki) on
response apple (Malus sp.), and European corn borer (Ostrinia
of insecta
nubilalis) on field corn (Luna, 1988). Again evidence
N, P, K M. persicae Tobacco +/∧/+
suggests that high levels of chemical fertilizer appli-
N Schizaphis graminum Oats/rye −
(greenbug) cations can cause nutritional imbalances in crops, in
N, lime S. graminum Oats − turn making them more susceptible to insect disease
N R. maidis Sorghum + pressure.
N, K, Ca M. persicae Brussels +/∨/− Because plants are a source of nutrients to herbiv-
sprouts
orous insects, an increase in the nutrient content of
N, P Therioaphis maculate Alfalfa −/+
(spotted alfalfa aphid) the plant maybe argued to increase its acceptability
a Symbols: (+) increase in density with increasing rates of as a food source to pest populations. Variations in
fertilizer element, (∧) highest density occurred at intermediate rates herbivore response may be explained by differences
of fertilizer element; (−) decrease in density with increasing rates in the feeding behavior of the herbivores themselves
of fertilizer element; (∨) lowest density occurred at intermediate (Pimentel and Warneke, 1989). For example, with in-
rates of fertilizer element. Slash separates the effects of fertilizer creasing N concentrations in creosotebush (Larrea tri-
elements listed in nutrients column.
dentate) plants, populations of sucking insects were
found to increase, but the number of chewing insects
populations associated with Brassica crop plants declined. With higher N fertilization, the amount of
have also been reported to increase in response to nutrients in the plant increases, as well as the amount
increased soil N levels (Table 3). In a 2-year study, of secondary compounds that may selectively affect
Brodbeck et al. (2001) found that populations of the herbivore feeding patterns. Protein digestion inhibitors
thrips Frankliniella occidentalis were significantly that accumulated in plant cell vacuoles were not con-
higher on tomatoes that received higher rates of N sumed by sucking herbivores, but inhibited chewing
fertilization. Seasonal trends in F. occidentalis on herbivores (Mattson, 1980).
tomato were found to be correlated to the number of In reviewing 50 years of research relating to crop
flowers per host plant, that changed with the N sta- nutrition and insect attack, Scriber (1984) found 135
tus of flowers. Plants subjected to higher fertilization studies showing increased damage and/or growth of
rates produced flowers that had higher N content as leaf-chewing insects or mites in N-fertilized crops,
well as variations in several amino acid profiles that versus fewer than 50 studies in which herbivore dam-
coincided with peak thrips population density. Abun- age was reduced by normal fertilization regimens.
dance of F. occidentalis (particularly adult females) In aggregate, these results suggest a hypothesis with
was most highly correlated to flower concentrations implications for fertilizer use patterns in agriculture,
of phenylalanine during population peaks. Other in- namely that high N inputs can precipitate high lev-

Table 3
Response of herbivores to increased soil nitrogen levels on Brassica host plants (Letourneau, 1988)
Host plant Herbivore species Factor Response

Brussels sprouts M. persicae No. of progeny Increase

Brussels sprouts B. brassicae No. of progeny Small increase, dependent on factors such as K
Rape Artogeia rapae Oviposition frequency Increase
Kale and cabbage A. rapae Oviposition frequency Increase
Kale A. rapae Oviposition frequency Increase
Cabbage A. rapae Growth rate Increase
Cabbage A. rapae Growth rate ultimate size Increase
Cabbage Plutella xylostella Feeding preference Increase
 M.A. Altieri, C.I. Nicholls / Soil & Tillage Research 72 (2003) 203–211 207

els of herbivore damage in crops. As a corollary, 4. The dynamics of insect herbivores in

crop plants would be expected to be less prone to organically managed systems
insect pests and diseases if organic soil amendments
were used, these generally resulting in lower N con- Studies documenting lower abundance of several
centrations in the plant tissue. Perhaps achieving insect herbivores in low-input systems have partly
more uniform foliar N concentration throughout the attributed such reductions to the lower N content in
year, avoiding pulse high foliar N levels following organically farmed crops (Lampkin, 1990). In Japan,
N fertilizer application, may be a key strategy to density of immigrants’ of the planthopper species
achieve optimum crop nutritional levels that deter pest Sogatella furcifera was significantly lower and the
attack. settling rate of female adults and survival rate of im-
Letourneau (1988) however questions if the mature stages of ensuing generations were generally
“N-damage” hypothesis, based on Scriber’s review, lower in organic compared to conventional rice fields.
can be extrapolated to a general warning about Consequently, the density of planthopper nymphs
fertilizer inputs associated to insect pest attack in and adults in the ensuing generations was found
agroecosystems. Of 100 studies of insects and mites to decrease in organically farmed fields (Kajimura,
on plants treated experimentally with high and low 1995).
N fertilizer levels, Letourneau found two-thirds of In England, conventional winter wheat fields exhib-
the studies to show an increase in insect growth, ited a larger infestation of the aphid Metopolophium
survival, reproductive rate, population densities or dirhodum than their organic counterpart (Kowalski
plant damage levels in response to increased N fer- and Visser, 1979). The conventionally fertilized wheat
tilizer. The remaining third of the arthropod studies crop had higher levels of free protein amino acids in
showed either a decrease in damage with fertilizer its leaves during June, which were attributed to a N
N or no significant change. The author noted, how- top dressing applied early in April. However, the dif-
ever, that experimental design can affect the types ference in the aphid infestations between crops was
of responses observed, which poses a problem for attributed to the aphid’s response to the relative pro-
insect responses to chemical and organic fertilization portions of certain non-protein to protein amino acids
treatments. present in the leaves at the time of aphid settling on
Firstly, the majority of the studies were conducted crops. The authors concluded that chemically fertil-
with potted plants versus less than 10% conducted in ized winter wheat was more palatable than its organ-
large-scale crop fields, which would have provided ically grown counterpart; hence the higher level of
a more realistic set of conditions for both plant N infestation.
uptake and subsequent herbivore response. Secondly, In greenhouse experiments, when given a choice
the studies conducted in fields did not clearly support of maize grown on organic versus chemically fertil-
the N-damage hypothesis. Although the sample size ized soils collected from nearby farms, European corn
was very small, the majority of comparisons showed borer (Ostrinia nubilalis) females significantly laid
no significant increase in arthropod performance or more eggs in the chemically fertilized plants (Phelan
damage with increased N. Even in field plot experi- et al., 1995). Interestingly, there was significant vari-
ments, the results were less than 60% in support of ation in egg-laying among chemical fertilizer treat-
the N-damage hypothesis. Only in greenhouse studies ments within the conventionally managed soil, but in
did the N-damage hypothesis hold true. Thirdly, ac- plants under the organic soil management, egg-laying
tual damage was measured in only 20% of the studies. was uniformly low. Pooling results across all three
Population levels (which could include different insect farms showed that variance in egg-laying was approx-
age classes) may be the next most important predictor imately 18 times higher among plants in convention-
of damage, but studies measuring these parameters ally managed soil than among plants grown under
were not found to support the N-damage hypothesis an organic regimen. The authors suggested that this
as much as those measuring parameters of growth, difference is evidence for a form of biological buffer-
survival, or reproductive rate in individual insect ing characteristically found more commonly in organ-
species. ically managed soils.
208 M.A. Altieri, C.I. Nicholls / Soil & Tillage Research 72 (2003) 203–211

Altieri et al. (1998) conducted a series of exper- techniques. Patzun farmers traditionally mixed ashes,
iments during 1989–1996 in which broccoli (Bras- kitchen scraps, crop residues, weeds, leaf litter, and
sica oleraceae) was subjected to varying fertiliza- manure to produce compost. However, since 1960,
tion regimes (conventional versus organic). The goal synthetic fertilizers were introduced to the region and
was to test the effects of different N sources on the were rapidly adopted in the area. Today, the majority
abundance of the key insect pests, cabbage aphid of farmers have replaced organic fertilizers with urea
(Brevicoryne brassicae) and flea beetle (Phyllotreta (CO(NH2 )2 ), although some recognize the negative
cruciferae). Conventionally fertilized monoculture consequences of the change and complain that pest
consistently developed a larger infestation of flea bee- populations have increased in their milpas since the
tles and in some cases of the cabbage aphid, than the introduction of the synthetic fertilizers.
organically fertilized broccoli systems. The reduction In their survey in the Guatemalan highlands,
in aphid and flea beetle infestations in the organically Morales et al. (2001) found that corn fields treated
fertilized plots was attributed to lower levels of free with organic fertilizer (applied for 2 years) hosted
N in the foliage of plants. This further supports the fewer aphids (Rhopalosiphum maidis) than corn
view that insect pest preference can be moderated by treated with synthetic fertilizer. This difference was
alterations to the type and amount of fertilizer used. attributed to a higher concentration of foliar N in corn
By contrast, a study comparing the population re- in the synthetic fertilizer plots, although numbers of
sponses of Brassica pests to organic versus synthetic fall armyworm (Spodoptera frugiperda) showed a
fertilizers, measured higher Phyllotreta flea beetles weak negative correlation with increased N levels.
populations on sludge-amended collard (B. oleracea) While fertilizers are under utilized in most parts of
plots early in the season compared to mineral-ferti- Asia, over-fertilization does occur in some regions,
lizer-amended and unfertilized plots (Culliney and especially where intensive vegetable production oc-
Pimentel, 1986). However, later in the season, in these curs. In addition to the cost, there are ecological and
same plots, insect population levels were lowest in or- health consequences of excessive fertilization (Con-
ganic plots for beetles, aphids and lepidopteran pests. way et al., 1991). Unused N from fertilizer can end
This suggests that the effects of fertilizer type varies up as nitrate in ground water, or in streams especially
with plant growth stage and that organic fertilizers where intensive vegetable crops are grown in high-
do not necessarily diminish pest populations but, at land areas (i.e. the Philippines, Thailand). A survey of
times, may unfortunately increase them. For example, 3000 dug wells in Indian villages showed that about
in a survey of California tomato producers, despite 20% of them contained nitrate levels in excess of the
the pronounced differences in plant quality (N content World Health Organization limit of 10 mg of NO3 -N
of leaflets and shoots) both within and among tomato per litter. Increased N levels have also been linked
fields, Letourneau et al. (1996) found no indication to increased pest problems in rice, notably the plant
that greater concentrations of tissue N in tomato plants brownhopper (Santikarm and Perkasem, 2000).
were associated with higher levels of insect damage.

6. Conclusions
5. Changes in pest status due to increased
fertilizer use Soil fertility management can have several effects
on plant quality, which in turn, can affect insect abun-
The majority of Cakchiquel farmers responding to dance and subsequent levels of herbivore damage. The
a survey conducted in Patzun, Guatemala, did not rec- reallocation of mineral amendments in crop plants can
ognize herbivorous insects as a problem in their corn influence oviposition, growth rates, survival and re-
milpas intercropped with beans, fava (Vicia fava), production in the insects that use these hosts (Jones,
and/or squash (Cucurbita maxima, C. pepo) (Morales 1976). Although more research is needed, preliminary
et al., 2001). The farmers attributed this lack of evidence suggests that fertilization practices can in-
pests to preventative measures incorporated into their fluence the relative resistance of agricultural crops to
agricultural practices, including soil management insect pests. Increased soluble N levels in plant tissue
 M.A. Altieri, C.I. Nicholls / Soil & Tillage Research 72 (2003) 203–211 209

following N fertilization, was found to generally de- can also provide supplies of secondary and trace el-
crease pest resistance, although this is not a universal ements, occasionally lacking in conventional farming
phenomenon (Phelan et al., 1995). systems that rely primarily on artificial sources of N,
Chemical fertilizers can dramatically influence the P, and K. Besides nutrient concentrations, optimum
balance of nutritional elements in plants, and it is fertilization, which provides a proper balance of ele-
likely that their excessive use will create nutrient im- ments, can stimulate resistance to insect attack (Luna,
balances, which in turn, reduce resistance to insect 1988). Organic N sources may allow greater tolerance
pests. Apparently N pulses following high fertilizer to vegetative damage in plants because such sources
applications leads to concentrations of foliar N which release N more slowly, during the course of one to
make plants more vulnerable to pest attack. In con- several years.
trast, organic farming practices, apparently promote Phelan et al. (1995) stressed the need to consider
an increase of soil organic matter and microbial activ- mechanisms other than N alone, when examining
ity and a gradual release of plant nutrients which does the link between fertility management and crop sus-
not lead to enhanced N levels in plant tissues, thus in ceptibility to insects. Their study demonstrated that
theory, allowing plants to derive a more balanced nutri- the ovipositional preference of a foliar pest can be
tion. Thus, while the amount of N immediately avail- mediated by differences in soil fertility manage-
able to the crop may be lower when organic fertilizers ment. Thus, the lower pest levels widely reported
are applied, the overall nutritional status of the crop in organic-farming systems may, in part, arise from
appears to be improved. Organic soil fertility practices plant–insect resistances mediated by biochemical

Fig. 2. An approach to agroecosystem health.

210 M.A. Altieri, C.I. Nicholls / Soil & Tillage Research 72 (2003) 203–211

or mineral-nutrient differences in crops under such (Homoptera: Delphacidae). Res. Popul. Ecol. 37, 219–
management practices. In fact, we feel such results 224.
provide interesting evidence to support the view that Klostermeyer, E.C., 1950. Effect of soil fertility on corn earworm
damage. J. Econ. Entomol. 43, 427–429.
the long-term management of soil organic matter can
Kowalski, R., Visser, P.E., 1979. Nitrogen in a crop–pest
lead to better plant resistance against insect pests. interaction: cereal aphids. In: Lee, J.A. (Ed.), Nitrogen as
Clearly more studies comparing pest populations on an Ecological Parameter. Blackwell Scientific Publications,
plants treated with synthetic versus organic fertiliz- Oxford, UK, pp. 67–74.
ers are needed. Understanding the underlying effects Lampkin, N., 1990. Organic Farming. Farming Press Books,
of organic fertilization on plant health may lead us Ipswitch, UK.
Letourneau, D.K., 1988. Soil management for pest control: a
to new and better IPM and ISFM program designs.
critical appraisal of the concepts. In: Proceedings of the
As we accumulate knowledge regarding the relation- Sixth International Science Conference of IFOAM on Global
ships between soil fertility and insect pest attack, we Perspectives on Agroecology and Sustainable Agricultural
will be better placed to convert conventional systems Systems, Santa Cruz, CA, pp. 581–587.
of crop production to those that incorporate agroeco- Letourneau, D.K., Drinkwater, L.E., Shennon, C., 1996. Effects of
soil management on crop nitrogen and insect damage in organic
logical strategies to optimize soil organic fertilization,
versus conventional tomato fields. Agric. Ecosyst. Environ. 57,
crop diversity management and more natural systems 174–187.
of pest regulation without incurring yield penalties Lockeretz, W., Shearer, G., Kohl, D.H., 1981. Organic farming in
(Fig. 2). the corn belt. Science 211, 540–546.
Luna, J.M., 1988. Influence of soil fertility practices on
agricultural pests. In: Proceedings of the Sixth International
References Science Conference of IFOAM on Global Perspectives on
Agroecology and Sustainable Agricultural Systems, Santa Cruz,
CA, pp. 589–600.
Adkisson, P.L., 1958. The influence of fertilizer applications on
Magdoff, F., van Es, H., 2000. Building Soils for Better Crops.
population of Heliothis zea and certain insect predators. J. Econ.
SARE, Washington, DC.
Entomol. 51, 144–149.
Mattson Jr., W.J., 1980. Herbivory in relation to plant nitrogen
Altieri, M.A., Nicholls, C.I., 1990. Biodiversity, ecosystem
content. Annu. Rev. Ecol. Syst. 11, 119–161.
function and insect pest management in agricultural systems.
In: Collins, W., Qualset, C.O. (Eds.), Biodiversity in McGuiness, H., 1993. Living Soils: Sustainable Alternatives
Agroecosystems. CRC Press, Boca Raton, pp. 69–84. to Chemical Fertilizers or Developing Countries. Consumers
Policy Institute, New York.
Altieri, M.A., Schmidt, L.L., Montalba, R., 1998. Assessing the
effects of agroecological soil management practices on broccoli Merrill, M.C., 1983. Eco-agriculture: a review of its history and
insect pest populations. Biodynamics 218, 23–26. philosophy. Biol. Agric. Hort. 1, 181–210.
Barker, A., 1975. Organic vs. inorganic nutrition and horticultural Meyer, G.A., 2000. Interactive effects of soil fertility and herbivory
crop quality. HortScience 10, 12–15. on Brassica nigra. Oikos 22, 433–441.
Brodbeck, B., Stavisky, J., Funderburk, J., Andersen, P., Olson, S., Morales, H., Perfecto, I., Ferguson, B., 2001. Traditional
2001. Flower nitrogen status and populations of Frankliniella fertilization and its effect on corn insect populations in the
occidentalis feeding on Lycopersicon esculentum. Entomol. Guatemalan highlands. Agric. Ecosyst. Environ. 84, 145–
Exp. Appl. 99, 165–172. 155.
Conway, G.R., Pretty, J., 1991. Unwelcome Harvest: Agriculture Oelhaf, R.C., 1978. Organic Agriculture. Halstead Press, New
and Pollution. Earthscan, London. York.
Culliney, T., Pimentel, D., 1986. Ecological effects of organic Painter, R.H., 1951. Insect Resistance in Crop Plants. University
agricultural practices in insect populations. Agric. Ecosyst. of Kansas Press, Lawrence, KS.
Environ. 15, 253–256. Phelan, P.L., Mason, J.F., Stinner, B.R., 1995. Soil fertility
Eggert, F.P., Kahrmann, C.L., 1984. Responses of three vegetable management and host preference by European corn borer,
crops to organic and inorganic nutrient sources. In: Organic Ostrinia nubilalis, on Zea mays: a comparison of organic and
Farming: Current Technology and its Role in Sustainable conventional chemical farming. Agric. Ecosyst. Environ. 56,
Agriculture. Pub. No. 46. American Society of Agronomy, 1–8.
Madison, WI, pp. 85–94. Pimentel, D., Warneke, A., 1989. Ecological effects of manure,
Jones, F.G.W., 1976. Pests, resistance, and fertilizers. In: sewage sludge and other organic wastes on arthropod
Proceedings of the 12th Colloquium of the International Potash populations. Agric. Zool. Rev. 3, 1–30.
Institute on Fertilizer Use and Plant Health, Bern, Switzerland. Santikarm, M.K., Perkasem, B., 2000. The Growth and
Kajimura, T., 1995. Effect of organic rice farming on planthoppers: Sustainability of Agriculture in Asia. Oxford University Press,
reproduction of white backed planthopper, Sogatella furcifera Oxford.
 M.A. Altieri, C.I. Nicholls / Soil & Tillage Research 72 (2003) 203–211 211

Schuphan, W., 1974. Nutritional value of crops as influenced by Slansky, F., Rodriguez, J.G., 1987. Nutritional Ecology of Insects,
organic and inorganic fertilizer treatments. Qual. Plant Foods Mites, Spiders and Related Invertebrates. Wiley, New York.
Hum. Nutr. 23, 333–358. van Emden, H.F., 1966. Studies on the relations of insect
Scriber, J.M., 1984. Nitrogen nutrition of plants and insect invasion. and host plant. III. A comparison of the reproduction
In: Hauck, R.D. (Ed.), Nitrogen in Crop Production. American of Brevicoryne brassicae and Myzus persicae (Hemiptera:
Society of Agronomy, Madison, WI. Aphididae) on Brussels sprout plants supplied with different
Slansky, F., 1990. Insect nutritional ecology as a basis for studying rates of nitrogen and potassium. Entomol. Exp. Appl. 9, 444–
host plant resistance. Florida Entomol. 73, 354–378. 460.