<?xml version="1.0" encoding="ISO-8859-1"?><article xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:xlink="http://www.w3.org/1999/xlink" xmlns:xsi="http://www.w3.org/2001/XMLSchema-instance">
<front>
<journal-meta>
<journal-id>0120-0488</journal-id>
<journal-title><![CDATA[Revista Colombiana de Entomología]]></journal-title>
<abbrev-journal-title><![CDATA[Rev. Colomb. Entomol.]]></abbrev-journal-title>
<issn>0120-0488</issn>
<publisher>
<publisher-name><![CDATA[Sociedad Colombiana de Entomología]]></publisher-name>
</publisher>
</journal-meta>
<article-meta>
<article-id>S0120-04882008000200001</article-id>
<title-group>
<article-title xml:lang="en"><![CDATA[Diversification practices: their effect on pest regulation and production]]></article-title>
<article-title xml:lang="es"><![CDATA[Prácticas de diversificación: sus efectos en la regulación de plagas y en producción]]></article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author">
<name>
<surname><![CDATA[POVEDA]]></surname>
<given-names><![CDATA[KATJA]]></given-names>
</name>
<xref ref-type="aff" rid="A01"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname><![CDATA[GÓMEZ]]></surname>
<given-names><![CDATA[MARÍA ISABEL]]></given-names>
</name>
<xref ref-type="aff" rid="A02"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname><![CDATA[MARTÍNEZ]]></surname>
<given-names><![CDATA[ELIANA]]></given-names>
</name>
<xref ref-type="aff" rid="A03"/>
</contrib>
</contrib-group>
<aff id="A01">
<institution><![CDATA[,University, Waldweg  ]]></institution>
<addr-line><![CDATA[Göttingen ]]></addr-line>
<country>Alemania</country>
</aff>
<aff id="A02">
<institution><![CDATA[,Universidad Nacional de Colombia. Facultad de Agronomia. ]]></institution>
<addr-line><![CDATA[Bogotá ]]></addr-line>
<country>Colombia</country>
</aff>
<aff id="A03">
<institution><![CDATA[,Universidad Nacional de Colombia. Departamento de Biología ]]></institution>
<addr-line><![CDATA[Bogotá ]]></addr-line>
<country>Colombia</country>
</aff>
<pub-date pub-type="pub">
<day>00</day>
<month>12</month>
<year>2008</year>
</pub-date>
<pub-date pub-type="epub">
<day>00</day>
<month>12</month>
<year>2008</year>
</pub-date>
<volume>34</volume>
<numero>2</numero>
<fpage>131</fpage>
<lpage>144</lpage>
<copyright-statement/>
<copyright-year/>
<self-uri xlink:href="http://www.scielo.org.co/scielo.php?script=sci_arttext&amp;pid=S0120-04882008000200001&amp;lng=en&amp;nrm=iso"></self-uri><self-uri xlink:href="http://www.scielo.org.co/scielo.php?script=sci_abstract&amp;pid=S0120-04882008000200001&amp;lng=en&amp;nrm=iso"></self-uri><self-uri xlink:href="http://www.scielo.org.co/scielo.php?script=sci_pdf&amp;pid=S0120-04882008000200001&amp;lng=en&amp;nrm=iso"></self-uri><abstract abstract-type="short" xml:lang="en"><p><![CDATA[: The interest to shift pest management strategies from the intensive use of agrochemicals to more sustainable and ecologically friendly practices has increased in recent years. One alternative to conventional farming systems is the implementation of diversification practices that increase diversity in- and around- the field to increase the incidence of natural enemies, reduce pest pressure and enhance crop production. In this review we illustrate the theoretical framework on which diversification practices are based and contrast it with the empirical evidence. The detailed review of 62 original studies published in the last ten years, shows that diversification practices (a) enhance natural enemies in 52%, (b) reduce pest pressure in 53% and (c) increase yield in only 32% of the cases where this was examined. We discuss these results on the basis of the reviewed studies providing key elements that should be taken into account to design diversification practices that can be implemented as competitive pest management strategies that cover the farmers’ needs, reducing the intensive use of agrochemicals.]]></p></abstract>
<abstract abstract-type="short" xml:lang="es"><p><![CDATA[El interés por dirigir las estrategias de manejo de plagas desde el uso intensivo de agroquímicos a prácticas sostenibles y ecológicamente amigables se ha incrementado en los últimos años. Una alternativa para los sistemas de cultivo convencionales es la diversificación tanto dentro como alrededor de los cultivos buscando incrementar la incidencia de enemigos naturales, reducir la presión de las plagas e incrementar o mantener la producción del cultivo. Se presenta una revisión del marco teórico que ha sido base para el estudio de las prácticas de diversificación y se contrasta con la evidencia empírica. Los resultados reportados en 62 estudios originales publicados en los últimos diez años, muestran que las prácticas de diversificación (a) incrementan los enemigos naturales en el 52% de los casos, (b) reducen la presión de las plagas en un 53% de los estudios e (c) incrementan los rendimientos en solo el 32% de los casos. Se discuten estos resultados teniendo como base los estudios que proveen elementos claves para ser tomados en cuenta para el diseño de prácticas de diversificación que puedan ser implementadas como estrategias competitivas de manejo de plagas y que cubran las necesidades de los productores reduciendo el uso intensivo de agroquímicos.]]></p></abstract>
<kwd-group>
<kwd lng="en"><![CDATA[Crop yield]]></kwd>
<kwd lng="en"><![CDATA[Intercrop]]></kwd>
<kwd lng="en"><![CDATA[Flowering-plant]]></kwd>
<kwd lng="en"><![CDATA[Repellent-plant]]></kwd>
<kwd lng="en"><![CDATA[Trap- plant]]></kwd>
<kwd lng="es"><![CDATA[Producción]]></kwd>
<kwd lng="es"><![CDATA[Policultivos]]></kwd>
<kwd lng="es"><![CDATA[Plantas con flores]]></kwd>
<kwd lng="es"><![CDATA[Plantas repelentes]]></kwd>
<kwd lng="es"><![CDATA[Plantas trampa]]></kwd>
</kwd-group>
</article-meta>
</front><body><![CDATA[ <p align="right"><font size="2" face="Verdana"><b>Art&iacute;culo de Revisi&oacute;n</b></font></p>     <p align="center">&nbsp;</p>     <p align="center"><font size="4" face="Verdana"><b>Diversification practices: their effect on pest regulation and production</b></font></p>     <p align="center">&nbsp;</p>     <p align="center"><font size="3" face="Verdana"><b>Pr&aacute;cticas de diversificaci&oacute;n: sus efectos en la regulaci&oacute;n de plagas y en producci&oacute;n</b></font></p>     <p>&nbsp;</p>     <p><font size="2" face="Verdana"><b>KATJA POVEDA<sup>1</sup>, MAR&Iacute;A ISABEL G&Oacute;MEZ<sup>2</sup> and ELIANA MART&Iacute;NEZ<sup>3</sup></b></font></p>     <p><font size="2" face="Verdana"><sup>1</sup> Postdoctoral researcher. Ph.D. Agroecology, G&ouml;ttingen University, Waldweg 26, 37073 G&ouml;ttingen, Alemania. <a href="mailto:kpoveda@gwdg.de.">kpoveda@gwdg.de.</a> Autor para correspondencia.</font></p>     <p> <font size="2" face="Verdana"><sup>2</sup> Researcher. MSc. Universidad Nacional de Colombia. Facultad de Agronomia. Ciudad Universitaria. Bogot&aacute;, Colombia. <a href="mailto:migomezj@unal.edu.co.">migomezj@unal.edu.co.</a></font></p>     <p> <font size="2" face="Verdana"><sup>3</sup> Researcher. MSc. Universidad Nacional de Colombia. Departamento de Biolog&iacute;a. Ciudad Universitaria. Bogot&aacute;, Colombia. <a href="mailto:emartinezpa@unal.edu.co.">emartinezpa@unal.edu.co. </a></font></p> <font size="2" face="Verdana">     ]]></body>
<body><![CDATA[<center> </center> </font> <hr size="1" />     <p><font size="2" face="Verdana"><b><font size="3">Abstract: </font></b>The interest to shift pest management strategies from the intensive use of agrochemicals to more sustainable and ecologically friendly practices has increased in recent years. One alternative to conventional farming systems is the implementation of diversification practices that increase diversity in- and around- the field to increase the incidence of natural enemies, reduce pest pressure and enhance crop production. In this review we illustrate the theoretical framework on which diversification practices are based and contrast it with the empirical evidence. The detailed review of 62 original studies published in the last ten years, shows that diversification practices (a) enhance natural enemies in 52%, (b) reduce pest pressure in 53% and (c) increase yield in only 32% of the cases where this was examined. We discuss these results on the basis of the reviewed studies providing key elements that should be taken into account to design diversification practices that can be implemented as competitive pest management strategies that cover the farmers&#8217; needs, reducing the intensive use of agrochemicals.</font></p>     <p>   <font size="2" face="Verdana"><b><font size="3">Key words: </font></b>Crop yield. Intercrop. Flowering-plant. Repellent-plant. Trap- plant.</font></p> <hr size="1" />     <p>   <font size="2" face="Verdana"><b><font size="3">Resumen:</font></b><font size="3"> </font>El inter&eacute;s por dirigir las estrategias de manejo de plagas desde el uso intensivo de agroqu&iacute;micos a pr&aacute;cticas sostenibles y ecol&oacute;gicamente amigables se ha incrementado en los &uacute;ltimos a&ntilde;os. Una alternativa para los sistemas de cultivo convencionales es la diversificaci&oacute;n tanto dentro como alrededor de los cultivos buscando incrementar la incidencia de enemigos naturales, reducir la presi&oacute;n de las plagas e incrementar o mantener la producci&oacute;n del cultivo. Se presenta una revisi&oacute;n del marco te&oacute;rico que ha sido base para el estudio de las pr&aacute;cticas de diversificaci&oacute;n y se contrasta con la evidencia emp&iacute;rica. Los resultados reportados en 62 estudios originales publicados en los &uacute;ltimos diez a&ntilde;os, muestran que las pr&aacute;cticas de diversificaci&oacute;n (a) incrementan los enemigos naturales en el 52% de los casos, (b) reducen la presi&oacute;n de las plagas en un 53% de los estudios e (c) incrementan los rendimientos en solo el 32% de los casos. Se discuten estos resultados teniendo como base los estudios que proveen elementos claves para ser tomados en cuenta para el dise&ntilde;o de pr&aacute;cticas de diversificaci&oacute;n que puedan ser implementadas como estrategias competitivas de manejo de plagas y que cubran las necesidades de los productores reduciendo el uso intensivo de agroqu&iacute;micos.</font></p>     <p> <font size="2" face="Verdana"><b><font size="3">Palabras clave:</font></b> Producci&oacute;n. Policultivos. Plantas con flores. Plantas repelentes. Plantas trampa.</font></p> <hr size="1" />     <p><font size="3" face="Verdana"><b> Introduction</b></font></p>     <p><font size="2" face="Verdana"> The use of chemically synthesized fertilizers and pesticides to reduce crop pests and weeds and to increase harvest yields is common in current agricultural practices. These practices are coupled with the removal of weeds from within and around crops, large field sizes, tillage operations of varying intensity and the degradation or destruction of non-crop habitats (reviewed by Gurr <i>et al</i>. 2003). Although these practices have substantially increased yield, they also increased production costs, pesticide resistance and have affected ecosystem and human health (Matson <i>et al</i>. 1997; Krebs <i>et al</i>. 1999; Tilman <i>et al</i>. 2002). At the ecosystem level they caused serious ecological problems such as water contamination, habitat degradation and loss of biodiversity (Matson <i>et al</i>. 1997 ; Krebs <i>et al</i>. 1999; Staver <i>et al</i>. 2001; Tilman <i>et al</i>. 2002) with the concomitant loss of ecological functions such as pollination and biological control (Kruess and Tscharntke 1994; Matthies and Schmid-Hempel 1995; Didham <i>et al</i>. 1996; Kruess and Tscharntke 2000; Tilman <i>et al</i>. 2002). In response to these negative effects, the world market has increased its demand for residue free food (Thompson 1998; Magnusson and Cranfield 2005). One alternative to conventional farming practices is the increase of in-and around- crop diversity to reduce pest pressure. It has been generally assumed that this practice stimulates the presence of natural enemies and enhances pest suppression, potentially reducing the need for costly and ecologically disruptive insecticide applications (i.e. Altieri and Nicholls 1994; Gurr <i>et al</i>. 2004). However, in order to propose technological packages that can be implemented by the farmers, the link between diversification practices and increased crop yield must be successfully shown (Gurr and Wratten 1999). There is an extensive theoretical literature predicting that biodiversity could enhance natural enemies and increase pest suppression (see next section). Also, empirical studies have tested the relationship between species diversity and functioning of natural enemy assemblages and pest suppression (Cardinale <i>et al</i>. 2003; Wilby and Thomas 2002a; Wilby and Thomas 2002b; Finke and Denno 2004; Straub et al. 2008), but a convincing link between habitat diversification, pest suppression and crop production seems to be missing.</font></p>     <p><font size="2" face="Verdana"> Our goal is to contrast the theoretical and empirical evidence on how diversification practices affect natural enemies, pest pressure and crop yield. We start this review by summarizing the theoretical background, then we review original literature to determine if the theoretical expectations are met in the empirical work, emphasizing studies done on crop yield. Based   on the results of this review we discuss the possible causes of   any discrepancy between theory and observation, and propose   some guidelines for future studies to develop management   practices that meet the needs of farmers and reduce the use of   agrochemicals.</font></p>     <p align="left"><font size="2" face="Verdana"><b>How does diversity increase pest control and   production? Theoretical background</b></font></p>     <p><font size="2" face="Verdana">   There have been several hypotheses to explain how vegetation   diversity can directly affect crop pests. In general, vegetation   diversity has been proposed to disrupt the pest&#8217;s ability to locate   the host plant, to increase mortality of the pest or to repel the   pest. Here we give a brief overview of the hypotheses that   have been proposed until now:</font></p>     ]]></body>
<body><![CDATA[<p><font size="2" face="Verdana">   a. <b>The disruptive crop hypothesis</b> is equivalent to Root&#8217;s   (1973) resource concentration hypothesis and stipulates that   herbivores in polycultures will have more difficulties finding   crop plants associated with one or more taxonomically or   genetically different plants than finding crop plants in   monoculture (Vandermeer 1989).</font></p>     <p><font size="2" face="Verdana">   b.<b> The trap crop hypothesis</b> suggests that pests will be   attracted to associated plants and hence are less likely to leave   the trap crop and wander into the principal crop (Vandermeer   1989).</font></p>     <p><font size="2" face="Verdana">   c. <b>The natural enemy hypothesis</b> proposes that a lower   number of phytophagous insects are found in complex   environments because predators and parasitoids are more   diverse and abundant in those environments compared to simple   environments (Root 1973; Russell 1989).</font></p>     <p><font size="2" face="Verdana">   d. <b>The barrier crop hypothesis</b> or physical obstruction   hypothesis bases its effectiveness on the use of taller non-host   plants to obstruct the movement of the pest insect within the   cropping system (Perrin and Phillips 1978).</font></p>     <p><font size="2" face="Verdana">   e.<b> The visual camouflage hypothesis</b> also known as the   &#8220;apparency hypothesis&#8221; incorporates the visual stimuli that   induce herbivores to land on plants: color and plant height.   Herbivores tend to land on tall green plants, so that using noncrop   plants to make the crop &#8220;less apparent&#8221; by adding more   green or taller plants is a useful mechanism to camouflage the   crop (reviewed by Finch and Collier 2000).</font></p>     <p><font size="2" face="Verdana">   f. <b>The associational resistance hypothesis</b> proposes that nonhost   plants confer protection to the crop by releasing &#8220;odor   masking&#8221; substances into the air making the crop &#8220;invisible&#8221;   to the herbivore (Tahvanainen and Root 1972).</font></p>     <p><font size="2" face="Verdana">   g. <b>The repellent chemicals hypothesis</b> predicts that the nonhost   plants emit odors that repel the herbivore (Uvah and Coaker   1984).</font></p>     <p><font size="2" face="Verdana">   h. <b>The altered profile of the host plant odor hypothesis</b> bases   its effect on changes in the physiology of the plant through   certain chemicals they take up from the soil (reviewed by Finch   and Collier 2000).</font></p>     <p><font size="2" face="Verdana">   The above hypotheses are supported in most cases by   experimental evidence (reviewed by Finch and Collier 2000).   However, the application of these techniques would be useless   for agriculture if pest suppression and enhanced natural enemies   do not translate into increased yield. Studies showing the link   between pest suppression and yield are limited (Ostman <i>et al</i>.   2003; Cardinale <i>et al</i>. 2003) but the results are promising.   Ostman <i>et al</i>. (2003) showed that ground-living natural enemies   (ground beetles, Carabidae; rove beetles, Staphylinidae and   spiders) of the bird cherry-oat aphid Rhopalosiphum padi L.,   1758 dramatically reduce aphid abundance. Aphid suppression   led to a 23% increase in barley Hordeum vulgare L. (Poaceae)   yield compared to scenarios where natural enemies of the bird   cherry-oat aphid were absent. In another study performed by   Cardinale <i>et al</i>. (2003) on the effect of three natural enemies   (Harmonia axyridis Pallas, 1773, Coccinellidae; Nabis sp.,   Nabidae and Aphidius ervi Haliday, 1834 Braconidae) on the   pea aphid Acyrthosiphon pisum Harris, 1776 (Aphididae) that   feeds on alfalfa Medicago sativa L. (Fabaceae) they found an   indirect effect of natural enemies on production mediated by   herbivore suppression. The presence of all three enemy species   reduced pea aphid density in the field. Crop yield was inversely   related to pea aphid density and therefore the presence of   natural enemies should increase yield. Although previous   studies seem to be very promising, we have to take into account   that those studies actively manipulate the presence of natural   enemies in the field (Cardinale <i>et al</i>. 2003; Ostman <i>et al</i>. 2003),   not reflecting what would happen in an agricultural setting.   Thus the question remains open if diversification practices   actually do increase the presence of natural enemies and   increase pest suppression as would be predicted from the above   hypotheses.</font></p>     <p><font size="2" face="Verdana"><b>   Effect of diversification on natural enemies, herbivores   and crop damage and production</b></font></p>     ]]></body>
<body><![CDATA[<p><font size="2" face="Verdana">   Theory predicts that diversified crops in and around the field   should have a higher and more effective population of natural   enemies, decreased pest pressure on the crop and consequently   higher yields in comparison to monoculture. In order to test   this prediction we searched for articles published in scientific   journals in the last ten years that investigated the effect of   diversification practices, like intercropping and local habitat   manipulation, on pest suppression and biological control. To   avoid biasing the articles with respect to known authors, groups   or papers, we searched the literature database (ISI Web of   Knowledge: http://isiknowledge.com) using the keywords:   &#8220;pest* AND diversification&#8221;, &#8220;pest* AND intercrop*&#8221;, &#8220;pest*   AND habitat manipulation&#8221;, &#8220;habitat manipulation AND   agroecosystems&#8221;, &#8220;biological control AND agroecosystems&#8221;,   &#8220;biological control AND habitat manipulation&#8221;. Out of the 279   references obtained in our search, we used the following criteria   to finally select the 62 references included in our analysis (Table   1): (1) studies should be conducted at a local scale, including   diversification practices in and immediately around the crop,   (2) the timing of crop growth and diversification practices   should be the same, excluding practices like crop rotation, (3)   only studies performed in the field and on crops or their   associated organisms are included, and (4) only studies that   were available to us through the online libraries of the   University of G&ouml;ttingen (Germany) and Cornell University   (USA) were included. For each study we recorded the crop,   the diversification mechanism used, the effects (positive,   negative and/or neutral) reported on herbivores, natural   enemies, crop damage and crop production. Diversification   practices were categorized into techniques performed in (52   studies) and around (seven studies) the crop and these   categories were further subdivided by the type of plant that   was used to increase diversity. Twenty-three studies increased   within field diversity with other crops (&#8220;in-crop&#8221;), seven studies   used flowering plants in the field (&#8220;in-flowers&#8221;) to attract natural   enemies and four studies used flowering plants around the crop   (&#8220;around-flowers&#8221;). The rest of the studies increased in-field   or around-field diversity by specific functional groups like trap-   (&#8220;in-trap&#8221;- five studies) or repellent- plants (&#8220;in-repellent&#8221;- four   studies) to attract or repel herbivores, by crops around the field   (&#8220;around-crop&#8221;- two studies) or non-specifically by using   weeds, ground cover plants, or natural diversity (&#8220;in-other&#8221;-   13 studies, and &#8220;around-other&#8221;- one study)  (<a href="img/revistas/rcen/v34n2/v34n2a01tab1.gif" target="_blank">Table 1</a>). Only   three studies used combined in- and around-field practices and   push-pull (&#8220;in/around-push-pull&#8221;) strategies to simultaneously   attract herbivores to trap plants around the field and repel   herbivores from the center of the crop (studies 34, 39, 40 in    (<a href="img/revistas/rcen/v34n2/v34n2a01tab1.gif" target="_blank">Table 1</a>)). In order to quantify if diversification practices   decreased, increased or had no effect on natural enemies,   herbivores and production, we independently scored each of   the effects reported in each study. In those cases where more   than one effect was shown, as for example in ), who reported   different effects on different species of natural enemies, we   scored each reported effect independently. Thus, we had 62   articles that report 171 effect cases. Studies reporting   contradictory effects on the same species in different locations   or in different years or on different ways of measuring the same   response, were quantified as an unclear response. For example,   Bukovinszky <i>et al</i>. (2004) reported a positive effect of   intercropping on the number of <i>Plutella xylostella</i> per broccoli   plant but a negative effect of the same treatment on the   abundance of P. xylostella at a plot level, so the effect on the   herbivore was scored as unclear.</font></p>     <p><font size="2" face="Verdana">   From the 62 studies only nine (studies 32-34, 41, 44, 47,   50, 54 &amp; 60) actually report positive effects of diversification   practices on yield coupled with enhanced presence of natural   enemies and / or a reduction in pest pressure. Eight studies   showed that diversification practices can cause a reduction in   yield as a consequence of both positive as well as negative   effects on herbivores, crop damage and / or natural enemies   (studies 8, 13, 26, 31, 43, 51, 55 &amp; 62). Most of the studies,   however, just reported effects on natural enemies, pest presence or yield and there is high variation in the effect of diversification on each of these response variables. Diversification effects on   natural enemy populations were recorded in 35 of the 62   studies. Natural enemies were quantified in terms of abundance   or (activity) density of parasitoids (study 18), predators (studies   10, 21, 23-25, 27, 28, 36, 37, 41, 42, 44, 46, 48 &amp; 49), or   natural enemies in general (studies 11, 14, 17, 19 &amp; 20), species   richness or diversity of predators (study 37), parasitism rates   (studies 3, 4, 6, 8, 12, 16, 23, 26, 34, 47 &amp; 50) or predation   rates (studies 3 &amp; 36). In those studies where natural enemies   were investigated, 52% of the cases reported a positive effect   of diversification practices. The diversification practices that   frequently led to an enhancement of natural enemies were   increasing abundance of flowering plants in the crop (studies   3, 6, 19, 29 &amp; 30), enhancing flowering plants around the crop   (studies 15, 48), intercropping mechanisms (studies 12, 14,   21, 53, 54, 60, 62), increasing in-field plant diversity nonspecifically   (25, 26, 46, 55), increasing in-field diversity with   repellent plants (52), and push-pull strategies (studies 39 &amp;   40) (<a href="img/revistas/rcen/v34n2/v34n2a01fig1.gif">Fig. 1A</a>). In 20% of the cases there was no effect of   diversification on natural enemies and only 9.5% of the cases   reported a negative effect of diversification. Unclear effects   were reported in 18.5% of the cases (<a href="img/revistas/rcen/v34n2/v34n2a01tab1.gif" target="_blank">Table 1</a>).</font></p>   <font size="2" face="Verdana">     <p>   In 44 out of the 62 articles the effect of diversification   practices on crop herbivores was quantified. Herbivores were   quantified in terms of larval infestation (study 2), number of   eggs (studies 7, 12 &amp; 59), density or abundance of immatures   and adults (studies 5, 8, 9, 14, 16-20, 22, 23, 25-29, 35-37, 41-   43, 45, 47, 50-53, 55-58, 60 &amp; 62), species richness (study 11)   and oviposition preferences (studies 39 &amp; 44). Overall, 53%   of the reported cases showed a negative effect of diversification   on herbivores as would be expected by theory. The   diversification practices that seem to be most effective in   leading to a herbivore reduction were intercropping (studies   8, 17, 21, 41, 42, 47, 50, 51, 53, 60 &amp; 62 reporting the expected   effects), non-specific in-crop diversity increase (studies 7, 16,   22, 26, 27, 44 &amp; 55 reporting the expected effects) and pushpull   strategies (studies 39 &amp; 40) (<a href="img/revistas/rcen/v34n2/v34n2a01fig1.gif">Fig. 1B</a>). </p>     <p> Diversification   practices had a positive effect on herbivore presence in 11.9%   of the analyzed cases, no effect in 22% of the cases and an   unclear effect in 13.1% of the cases (<a href="img/revistas/rcen/v34n2/v34n2a01tab1.gif" target="_blank">Table 1</a>).   Effects on plant damage were reported in 18 studies and   were quantified in terms of foliage consumption (studies 8, 50   &amp; 51), deposits of frass and tunneling (study 28), tissue damage   (studies 9, 16, 35, 38, 54 &amp; 62), stem boring (studies 12, 23,   33, 34, 44 &amp; 62), root knotting (study 31) or root necrosis   (study 38). In 57.9% of the reported cases, plant damage was   reduced with diversification practices. This expected effect was   achieved when implementing diversification practices like   intercropping (studies 8, 13, 51, 54 &amp; 62), in-field use of   repellent plants (studies 32 &amp; 33), push-pull practices (study   34) and the use of non-specific plants in (studies 26, 28 &amp; 44)   and around the crop (study 23) (<a href="img/revistas/rcen/v34n2/v34n2a01fig1.gif">Fig. 1C</a>). Diversification   practices increased crop damage in 21.1% of the reported cases.   No effect of diversification practices on crop damage occurred   in 15.8% of the cases, while only 5.2% reported an unclear   effect (<a href="img/revistas/rcen/v34n2/v34n2a01tab1.gif" target="_blank">Table 1</a>).</p>     <p>   Effects on production were quantified in 30 of the 62   studies. Production was quantified in terms of yield (studies 1,   9, 13, 19, 22, 24, 26, 32-35, 38, 41, 43-45, 47, 49, 50-56, 60 &amp;   62), size of the product (studies 8 &amp; 27) and development time   (study 31). There was a positive effect of diversification   practices in 32% of the cases. Out of all the diversification   mechanisms, the push-pull strategy reported a consistently   positive effect on production, however, this strategy was   represented by only one study evaluating effects on production   (study 34) (<a href="img/revistas/rcen/v34n2/v34n2a01fig1.gif">Fig. 1D</a>). Diversification practices had a negative   effect on production in 28.9% of the cases, no effects in 26.1%   of the cases and an unclear effect in 13% of the cases.</p>     <p><b>   <font size="3">Discussion</font></b></p>     <p>   We did not find that diversification practices consistently   enhance natural enemies, decrease herbivores, or increase   production. Rather, for natural enemies and herbivores, only   about half of the cases report the expected effects. Of even   more concern, for practicing farmers, only one third of the   cases report an increase in production. Given the somewhat   discouraging results we discuss the possible causes that lead   to these unexpected effects. Using studies that show positive   results as examples, we explore how particular approaches   could help future diversification studies achieve the expected   goals that will result in farmer adoption of these kinds of   technologies.</p>     <p><b>   The importance of the &#8220;right kind&#8221; of diversity</b>. Although   diversification practices base their effectiveness on the fact   that high diversity should lead to pest suppression, it is also   known that high plant diversity in agroecosystems does not   automatically reduce pest pressure and enhance the activity   of natural enemies (Landis <i>et al</i>. 2000; Heemsbergen <i>et al</i>.   2004). Several authors have noted that to selectively enhance   natural enemies, the functionally important elements of   diversity should be identified and provided, rather than   encouraging diversity per se (Landis <i>et al</i>. 2000). Heemsbergen   <i>et al</i>. (2004) suggest that it is not the species number but the   degree of functional differences between species that enhance   overall ecological functions. The species-specific contribution   to the range of functional groups in a community might be an   important mechanism by which biodiversity generates positive   interactions that enhance ecological services like pest   suppression. Therefore, the screening of key plants is of crucial   importance to shape agricultural systems to specifically reduce   pest pressure and enhance production.</p>     <p>   <b>Increasing diversity with other crops and plants</b>. The use of   other crops to reduce pest pressure and increase yield of the   main crop, known as intercropping, is a long established practice   (Vandermeer 1989; Altieri and Nicholls 1994). The effectiveness   of this practice is exemplified in one of the reviewed studies   where cowpea Vigna unguiculata (L.) Walp. (Fabaceae) and   okra Abelmoschus esculentus (L.) Moench (Malvaceae) were   intercropped with tomato Solanum lycopersicum L.   (Solanaceae) (Pitan and Olatunde 2006). Intercropping had a   negative effect on the herbivores and a positive effect on yield   in both crops, though the exact mechanism remains unclear. </p>     <p>  However, increasing diversity can increase pest problems   (reviewed by Landis <i>et al</i>. 2000). This undesired effect can be   avoided with knowledge of pest natural history. This was   certainly shown in the study by Ngeve (2003) where intercropping   cassava Manihot esculenta Crantz (Euphorbiaceae)   with maize Zea mays L. (Poaceae) and groundnuts Arachis   villosulicarpa Hoehne (Fabaceae) actually increased the   severity of root mealybug Stictococcus vayssierei Richard   (Stictococcidae) infestation. This increased pest pressure was   a consequence of using other mealybug host plants as the   intercropping species (Ngeve 2003). It becomes obvious from this example that knowledge of the alternative hosts of the   pest is crucial, in order not to add additional food resources to   a pest that is meant to be controlled. This factor is also important   when choosing flowering plants to attract natural enemies,   and will be discussed in the next section.</p> </font>    ]]></body>
<body><![CDATA[<p><font size="2" face="Verdana">   Regardless of the previously published work on how pest   suppression leads to an increased yield (Cardinale <i>et al</i>. 2003;   Ostman <i>et al</i>. 2003), our literature review demonstrates that   diversification practices that reduce pest pressure do not   necessarily achieve an increased production (i.e. Showler and   Greenberg 2003; Sastawa <i>et al</i>. 2004; Schulthess <i>et al</i>. 2004).   Mechanisms like competition and allelopathic effects between   plants could be responsible for these effects. Sastawa <i>et al</i>.   (2004) compared intercropping systems varying in their   complexity: simple intercrops of millet Pennisetum glaucum   (L.) R. Br. (Poaceae) and soybean <i>Glycine max</i> (L.) Merr.  (Fabaceae), and more complex intercrops of millet, soybean,   groundnut and cowpea. They found that the more complex   systems actually led to a reduction in the number of the pod   sucking bug<i> Nezara viridula</i> Linnaeus, 1758 (Pentatomidae)   and a reduction in the defoliation caused by two carabids   (<i>Egadroma discriminatum</i> Basi and <i>Siderodactylus sagitarius</i>  Meigen) to soybean. However, soybean yield also decreased   in the more complex diversification systems. The authors   suggest that competition and shading by the intercropped plants   were the possible causes for the reduced production (Sastawa   <i>et al</i>. 2004). Very similar results are reported by Schulthess et   al. (2004) and Showler and Greenberg (2003) where   diversification practices suppress the pest but simultaneously   reduce yield, probably as a consequence of competition. Moreover, empirical evidence shows that competition not only   decreases yield, but could also be the cause of reduced pest   pressure. Bukovinszky <i>et al</i>. (2004) assessed the effect of   intercropping Brussels sprouts Brassica oleracea var.   gemmifera D. C. (Brassicaceae) with malting barley (<i>H.   vulgare</i>) on the populations of P. <i>xylostella</i> and <i>Brevicoryne brassicae</i> L., 1758 (Aphididae). They reported a lower   incidence of both herbivores on the intercropped Brussels   sprout in comparison to monocrops, but the effect seemed to   be caused by the effect of competition between both plants.   Competition caused drought stress on Brussels sprout plants,   leading to reduced size and delayed phenology, which made   those plants less apparent and less attractive to the herbivore   (Bukovinszky <i>et al</i>. 2004). Effects of plant-plant interactions   like competition and allelopathy (Kamunya <i>et al</i>. 2008) can   negatively affect production and override positive effects on   pest suppression. The previous examples make clear that effects   on pest pressure cannot be simply extrapolated to crop yield   and that great caution has to be taken when choosing the plant   to intercrop.</font></p> <font size="2" face="Verdana">    <p>   The goal of diversifying crops is often to increase the   availability of appropriate microhabitats for the natural enemies   of the pests (Sunderland and Samu 2000; Gurr <i>et al</i>. 2003).   Examples from our literature review show that broccoli   (<i>Brassica oleracea var. botrytis</i> L. - Brassicaceae) stands intercropped   with different kinds of clover (<i>Trifolium fragiferum </i>L., <i>Trifolium repens</i> L., <i>Melilotus officinalis</i> (L.) Lam.) have   increased spider density and increased yield in comparison to   broccoli monocrops (Hooks and Johnson 2004). Also   intercropping maize with groundnut, soybean and Phaseolus   beans increases nesting of predatory ants in the field, reducing   termite attack and increasing yield (Sekamatte <i>et al</i>. 2003). In   both cases the enhanced predator presence is explained by the   provision of extra food resources and refuges as proposed by   Root (1973), making these desirable characteristics in the plants   used to intercrop. However, there is a confounding effect in   the last two studies when reporting a yield increase given by   the use of legumes as intercrop. Legumes are known for their   nitrogen fixing capacity that should increase the nitrogen   available to the main crop through organic residues and the   residual effect of the biologically fixed nitrogen (Lal <i>et al</i>.   1978). Although in the previous examples it is not clear if the   increased yield was accomplished by pest suppression or by   the presence of legumes, the desired effect of increased yield   was reached. The previous examples show that legumes are   excellent candidates for intercropping giving their   characteristics of enhancing the presence of natural enemies   and at the same time increasing yield. However, factors like   competition for resources can also be playing a role when   intercropping legumes. In one study Rao and Mathuva (2000)   report two different outcomes of intercropping legumes. They   showed that intercropping maize with pigeonpea Cajanus cajan   (L.) Millsp. (Fabaceae) increased yield by 24% in comparison   to monocultured maize, while intercropping maize with the   perennial legume Gliricidia sepium (Jacq. Kunth ex Walp.)   did not affect maize yield. The difference in the response was   attributed to the type of legume. The competition for water   between the superficial roots of Gliricidia and maize seem to   be the reason that there was no yield increase (Govindarajan   <i>et al</i>. 1996; Rao and Mathuva 2000). Negative yield effects as   a result of intercropping with a legume are reported by Harvey   and Eubanks (2004), who intercropped white clover (T. repens)   in broccoli to control P. xylostella with fire ants. Competition   lead to smaller, fewer and deformed broccoli leaves and finally   to a reduced yield. These latter studies show that although   legumes can have the added advantage of increasing yield   through their nitrogen fixing capacities, this effect cannot be   generalized for all legumes in all crops. Competition between   the chosen legume and the crop has to be tested before   implementing them in a diversification practice.</p>     <p>   Like plants from other groups, legumes can also have an   effect on pest oviposition. Bjorkman <i>et al</i>. (2007) showed that   the turnip root fly <i>Delia radicum</i> L., 1758 (Anthomyiidae)   reduced oviposition by approximately 50% when intercropping   cabbage <i>Brassica oleraceae</i> L. (Brassicaceae) with red clover   <i>T. pratense. </i>A similar result was reported by Chabi-Olaye <i>et   al</i>. (2005a) who showed that intercropping maize with legumes   could reduce the percentage of plants with stem borer eggs   also by approximately 50%. The incidence of <i>Thrips tabaci </i>Lindeman, 1889 (Thripidae) is also reduced when intercropping   leek <i>Allium porrum</i> L. (Liliaceae) with the legume <i>T. fragiferum</i>  (den Belder <i>et al</i>. 2000). In neither study was the effect on   production reported, thus it remains unclear if the negative   effect on herbivore oviposition translates into a positive effect   on production. Although none of the studies emphasized the   mechanism underlying the herbivore response, the disruption   of host finding could be a feasible explanation (Chabi-Olaye   <i>et al</i>. 2005a; Bjorkman <i>et al</i>. 2007), and changes in plant quality   through intercropping seem also to be playing a role (den Belder   <i>et al</i>. 2000).</p> </font>     <p>   <font size="2" face="Verdana"><b>Flowering plants to enhance natural enemies</b>. Potential   mechanisms of positive diversity effects include improving   the availability of alternative foods such as nectar, pollen and   honeydew for the natural enemies of pests (Patt <i>et al</i>. 1997;   Landis <i>et al</i>. 2000; Tylianakis <i>et al</i>. 2004). However, the mere presence of flowering plants in an agroecosystem is not always   sufficient to guarantee nectar supply for parasitoids (Baggen   and Gurr 1998; W&auml;ckers 2004) and identification of the key   flowering plants for certain parasitoids is required to guarantee   the enhancement of natural enemies. The first important factor   is to determine plant identity. Colley and Luna (2000) studied   the effect of 11 different flowering plants on the presence of   aphidophagous hoverflies (Syrphidae) giving an example of   how a screening process for a flowering plant takes place. However, it is important to take into account that resources   that are available for natural enemies could also be a food   source for herbivorous pests (Lavandero <i>et al</i>. 2006). For   example, Jones and Gillett (2005) intercropped polycultures with   sunflowers <i>Helianthus annuus </i>L. (Asteraceae), which increased   the presence of arthropod natural enemies (Jones and Gillett   2005) and insectivorous birds (Jones and Sieving 2006), but at   the same time herbivorous pests (Jones and Gillett 2005). For   this reason screening for suitable flowering plants should also   include the screening of the suitability for pest herbivores as   was done by Begum <i>et al</i>. (2006). They screened five flowering   plants to detect their effect on natural enemies and herbivores. </font></p> <font size="2" face="Verdana">    <p>   After greenhouse and field experiments they determined that     <i>Lobularia maritima</i> (L.) Desv. (Brassicaceae) provided benefits   to the egg parasitoid Trichogramma carverae Oatman and Pinto   (Trichogrammatidae) when mass released in vineyards, but not   on the leafroller pest <i>Epiphyas postvittana</i> (Walker)   (Tortricidae). Another important factor is that field conditions   and the type of management can alter the outcome of   diversification practices. Although the results from Begum et   al. (2006) seem very promising, the applicability to different   conditions seems to be inconsistent. Bell <i>et al</i>. (2006) used the   same species (L. <i>maritima</i>) in vineyards to control the same   type of pest (E. <i>postvittana</i>) but they did not find the same   results; plots intercropped with the flowering species did not   have increased parasitism rates. In this case biotic factors like   proximity to an orchard, which seems to be the source for   parasitoids, had a higher effect on parasitism than the increased   availability of local resources like L.<i> maritima</i>. This emphasizes   screening for the right flowering plant is not sufficient to   achieve the expected results, but that results from laboratory   settings or given field conditions may not yield the same effects   under different conditions.</p>     <p>   Using flowering plants around the crop could have the   disadvantage that the population of predators and parasitoids   stays within the flowering strips around the crop and does not   migrate to the field when resources in that strip are more   abundant (Rand <i>et al</i>. 2006). This was exemplified by the study   of Frere <i>et al</i>. (2007) where rose <i>Rosa rugosa </i>Thunb. (Rosaceae)   bushes were used to increase diversity around wheat   <i>Triticum aestivum</i> L. (Poaceae) fields. However, the presence   of rose bushes did not influence the aphid population within   the field. One likely explanation is the relatively higher   availability of resources such as pollen, nectar, aphid hosts for   predators and parasitoids in the rose borders.</p>     <p>   Although reviews and original studies (Baggen and Gurr   1998; Gurr and Wratten 1999; Landis <i>et al</i>. 2000; W&auml;ckers   2004; Lavandero <i>et al</i>. 2006) have already highlighted the   importance of selecting the appropriate flowering plant, our   literature review reveals that the link between enhancing natural   enemies through flowering plants and increasing crop yield is   still missing. Only one of the eleven studies in which diversity   was increased with flowering plants reported an effect on   production. Fitzgerald and Solomon (2004) found no effect on   apple Malus domestica Borkh. (Rosaceae) yield when the trees   were undersown with flowering plants. However, there is   evidence from other studies that flowering plants can reduce   yield, probably as a result of competition (Brown and Glenn   1999).</p>     <p>   <b>Repellent plants for herbivores.</b> An alternative method to   reduce pest pressure is to identify key plants that repel   herbivores (Vanhuis 1991; Finch <i>et al</i>. 2003; Lapointe <i>et al</i>.   2003; Morley <i>et al</i>. 2005). In this review only four studies that   used repellent plants against herbivores also studied their   effects on production. Two out of the four studies successfully   achieved the goal of reducing pests, increase yield and even   suppress weeds (Khan <i>et al</i>. 2006a; Khan <i>et al</i>. 2006b). One of   the studies (Khan <i>et al</i>. 2006b) exemplifies the importance of   continuing screening for appropriate plants, to cover the   different needs and the heterogeneity found in different   regions. Knowing that <i><i><i>Desmodium </i></i>uncinatum</i> (Jacq.) DC.   (Fabaceae) had the potential to control the stemborers Chilo   partellus (Swinhoe) (Crambidae) and <i>Busseola fusca</i> (F&uuml;ller)   (Noctuidae) on maize and suppress the witchweed Striga   <i>hermonthica</i> (Del.) Benth. (Scrophulariaceae), Khan <i>et al</i>.   (2006b) continued searching for the effectiveness of four other   species of <i><i>Desmodium </i></i>to be used under different agroecological   conditions. All <i><i>Desmodium </i></i>species tested achieved the same   results on stemborer suppression, witchweed control, and   maize yield increase as D. uncinatum. This result is the basis   for a technological tool that does not depend on a single   species, increasing the range of sites where the technology   can be implemented. Once a promising plant is identified as   having a repellent effect, its properties to control herbivores in   different crops should be investigated. This was performed by   Khan <i>et al</i>. (2006a), who studied the effectiveness of D.   uncinatum in sorghum <i>Sorghum bicolor</i> (L.) Moench (Poaceae)   fields after demonstrating their effectiveness in maize. They   found that with the same repellent plant (<i>D. uncinatum</i>) they   could achieve pest reduction, weed control, and increased yield   not only in maize but also in sorghum (Khan <i>et al</i>. 2006a),   increasing the applicability of a given technology to more than   one crop.</p>     <p>   Schader <i>et al</i>. (2005) reported that intercropping cotton   <i>Gossypium barbadense</i> L.(Malvaceae) with basil <i>Ocimum   basilicum</i> L. (Lamiaceae) as a repellent plant reduced pest   infestation and increased the abundance of the epigeic fauna.   However, no correlation between pest infestation and cotton   yield was detected; there was no decreased cotton yield even   though there was a 33% decrease in the amount of cotton   cultivated due to the intercropping. It is assumed that both a   basil-induced repellence against pest insects and a stimulation   of beneficial epigeic fauna might be responsible for the lower   pest infestation observed in intercropped plots.</p>     <p>   The previous results emphasize that the identification of   appropriate plants is a long-lasting process that is based on the   screening of hundreds of species (as will be discussed in the   section of push-pull strategies) or a longer history of research   on each plant. Moreover, it is very important to study the   chemical properties of plants such as repellent plants, to better   understand their interaction with the crop and pest, and to   permit future manipulation of the desired effects. For example   the reduced infestation by stemborers in maize&#8211;D. uncinatum   intercrops has been shown to be mediated by specific volatiles   released by the repellent plant (Khan <i>et al</i>. 2000). Knowing the   chemical properties of repellence not only permits a better understanding of the mechanisms, but it also gives the   possibility to produce synthetic volatiles, to simulate those of   the plant and have the potential to repel the herbivore or recruit   natural enemies (Pickett <i>et al</i>. 1997; Khan <i>et al</i>. 2008a). Using   molecular tools it may also be possible to modify the secondary   metabolism of the plant to release a higher concentration of   the repellent volatiles at all or only some stages of development   (Khan <i>et al</i>. 2008a).</p>     ]]></body>
<body><![CDATA[<p>   Not all pests react in the same way to repellent plants; what   can be very effective for one pest is not necessarily effective   on another pest. This was exemplified by the study of McIntyre   <i>et al</i>. (2001), who intercropped banana with three leguminous   crops, that had previously been reported as having repellent or   insecticidal properties on different pest species of different   crops. They failed to detect any negative effects of legumes on   the banana weevil<i> Cosmopolites sordidus </i>(Germar) (Curculionidae)   population and the presence of the nematodes   <i>Radopholus similis</i> (Cobb) and <i>Helicotylenchus</i> spp,   demonstrating that the repellence of several different organisms   does not mean that a plant will be effective on other pests.</p> </font>    <p>   <font size="2" face="Verdana"><b>Trap plants to attract herbivores</b>. Trap crops can be plants   of a preferred growth stage, cultivar, variety, or species that   are more attractive to the pest than the main crop. Thus trap   crops reduce herbivore pressure and concentrate the pest   population to a limited area, where it can be easily controlled   by traditional methods (Hokkanen 1991; Asman 2002; Shelton   and Nault 2004; Shelton and Badenes-Perez 2006). In this   literature review five studies used trap plants as intercrops to   control pests (Bender <i>et al</i>. 1999; Smith <i>et al</i>. 2000; Smith and   McSorley 2000; Badenes-Perez <i>et al</i>. 2005; Bullas-Appleton   <i>et al</i>. 2005). As for the repellent plants, the effective   identification and use of trap plants will depend on an   exhaustive screening of the potential trap crop (Khan <i>et al</i>.   2000), its effectiveness when using different crops or different   pests and the importance of local differences in abiotic and   biotic factors (Khan <i>et al</i>. 2008b). For example, Bender <i>et al</i>.   (1999) used Indian mustard <i>Brassica juncea</i> (L.) Czern.   intercropped in cabbage to study its effectiveness on   controlling lepidopterous larvae, mainly of the diamondback   moth <i>P. xylostella</i>. In the introduction of their study they   already report contradictory results of the effectiveness of   this potential trap species on the diamondback moth in   cabbage in regions as different as Taiwan, India, and Hawaii.   They tested the effectiveness of this trap species in Texas   and concluded that there was no effect of intercropping   cabbage with Indian mustard on any lepidopterous larvae. The actual causes of the differences achieved using the same   trap plant in the same crop on the same pest remains   inconclusive. However, it is clear that regional differences in   biotic or abiotic factors could determine the effectiveness of   such a practice. A similar case is reported in the paper by   Smith and McSorley (2000) who studied the effect of   intercropping eggplant <i>Solanum melongena</i> L. (Solanaceae)   as a trap crop for management of whiteflies <i>Bemisia argentifolii </i>Bellows &amp; Perring (Aleyrodidae) on bean <i>Phaseolus   vulgaris </i>L. (Fabaceae). They report no effect of the eggplant   intercropping system on the density of eggs and nymphs.   This experiment exemplifies that the trap plant used was not   effective under their growing conditions and they report that   air currents determine the migration of adult whiteflies into   plots, showing again that abiotic factors can be playing a crucial   role.</font></p> <font size="2" face="Verdana">    <p>   The importance of determining if a reduced pest pressure   translates into an increased productivity is a concern in the   studies with trap crops. Only one study showed the effect of   an attractive plant on pest suppression and production. Bullas-   Appleton <i>et al</i>. (2005) investigated the effect of inter-planting   the highly susceptible cultivar Berna Dutch brown bean as a   trap crop in the moderately susceptible cultivar Stingray white   bean<i> P. vulgaris </i>on pest pressure and yield. Although they   reported that at the beginning of the season intercropping   reduced damage on the plants by potato leafhoppers <i>Empoasca   fabae </i>(Harris) (Cicadellidae), this effect disappeared at the   end of the season, and there was no effect of intercropping   with trap plants on yield.</p> </font>    <p>   <font size="2" face="Verdana"><b>The integrated use of repellent and attractive plant stimuli:   the push-pull strategy</b>. From the previous section we could   infer that repellent stimuli seem to be very effective to reduce   pest pressure and increase yield, while trap plants seem not to   be as effective and their effects on production remain unclear. One possible reason for the mixed results when using trap   plants is that the local attraction sought in trap crops also   causes a regional attraction that increases the presence of the   pest in the field since they are more attracted from outside the   field by the trap plants (Vandermeer 1989). This negative effect   could be compensated for by the integrated use of behaviormodifying   stimuli to manipulate the distribution and abundance   of pests, which has been named a &#8220;push-pull&#8221; strategy. This   strategy is based on selectively increasing plant diversity to   decrease pest pressure by identifying key plants that repel   herbivores to make the protected culture unattractive for the   pests (push) (Vanhuis 1991; Lapointe <i>et al</i>. 2003), while at the   same time using trap plants that lure the pest toward them   (pull) (Hokkanen 1991). A review on the principles of this   strategy and the current knowledge is presented by Cook et   al. (2007).</font></p> <font size="2" face="Verdana">    <p>   Only three studies in our literature review evaluated the   effect of push-pull strategies as pest management systems   (Midega <i>et al</i>. 2006; Khan <i>et al</i>. 2008b; Midega <i>et al</i>. 2008).   All three studies were performed by the same group of   investigators and are based on the same system. They developed   a push-pull strategy to control the corn stemborers C. <i>partellus</i>  and <i>B. fusca</i> in maize fields from Kenya. This strategy is based   on the use of herbaceous plants of economic importance. The   push stimulus is an intercrop of the forage legume D.   <i>uncinatum</i>, and border rows of Napier grass <i>Pennisetum   purpureum</i> Schumach. (Poaceae) exert the pull effect. This   practice enhanced the abundance of natural enemies like spiders   (Midega <i>et al</i>. 2008), increased predation rates of <i>C. partellus </i>(Midega <i>et al</i>. 2006) and reduced oviposition of <i>C. partellus</i>  (Midega <i>et al</i>. 2006). Khan <i>et al</i>. (2008b) evaluated the   effectiveness of this attractive-repellent practice under farmers&#8217;   conditions, comparing the push-pull technology against maize   monocrops in 280 farms. Field surveys agree with the farmers&#8217;   perception that the push-pull strategy reduced stemborers and   increased yield. Besides controlling the stem borers and   increasing yield, witchweed (which decreases maize yield) is   also controlled (Khan <i>et al</i>. 2008b). Although the push-pull   technology seems to be achieving more than the expected   results of a diversification practice on pest suppression and   yield increase, we are aware that these results were only   obtained as a consequence of many years of studying the   system and its effectiveness (as can be inferred from the following studies: Khan <i>et al</i>. 1997; Khan <i>et al</i>. 2000; Khan   and Pickett 2004; Khan <i>et al</i>. 2006b; Cook <i>et al</i>. 2007). The   starting point to develop this technology involved a screening   process of several hundred plant species, mainly of the family   Poaceae, but also Cyperaceae, Thyphaceae and some   Fabaceae (Khan <i>et al</i>. 2000). The attack rate by the different   species of stem borers was examined and the colonization   rate was taken to choose potential trap plants (as being those   species with the highest colonization rates) and potential   repellent plants (as being the least attractive plants). The two   most attractive crop plants were Napier grass and Sudan   grass, Sorghum sudanensis Stapf (Poaceae), while the most   repellent plants were molasses grass, Melinis minutiflora   Beauv. (Poaceae) and two legume species, silverleaf, D.   uncinatum, and greenleaf, D. intortum (Mill.) Urb. (Fabaceae)   (Khan <i>et al</i>. 2000). The legumes had the added advantage of   suppressing development of the problematic weed S.   hermonthica. With these potentially effective trap and   repellent plants, experiments where performed in 1996. Napier   grass was highly effective as a trap plant since it attracted   most of the oviposition but at the same time reduced larval   survival on the plant to 20% (in comparison with 80% on   maize) (Khan <i>et al</i>. 2000). The effect was caused by the   production of sticky sap by the Napier grass that trapped   and killed the larvae (Khan and Pickett 2004). This effect was   confirmed in further years of experiments that showed a yield   improvement of more than 1 t/ha (Khan <i>et al</i>. 2000). The   effectiveness of intercropping with the repellent plants was   also confirmed in the field showing that the use of M.   minutiflora and <i><i>Desmodium </i></i>significantly reduced the   presence of the stemborers. The rate at which the repellent   plants had to be intercropped in the fields was also assessed   in further studies determining that <i>M. minutiflora</i> would be   ideally planted at a density of 1:3 although it could be planted   in densities of 1:10 while still achieving the expected results   (Khan <i>et al</i>. 2000). After choosing the plants responsible for   pest control, the mechanisms behind the effect were analyzed   to increase the robustness and reliability of this pest control   method. Plants use indirect defenses such as volatile organic   compounds (VOCs) to attract or repel herbivores and their   natural enemies (Karban and Baldwin 1997). Khan <i>et al</i>.   (2008a) reported that for stem borer control, the plant   chemistry responsible involves release of attractant VOCs   (hexanal, (E)-2-hexenal, (Z)-3-hexen-1-ol, (Z)-3-hexen-yl acetate)   from the trap plants and repellent VOCs ((E)-ocimene, &szlig;-   terpinolene, &szlig;-caryophyllene, humulene, (E)-4,8-dimethyl-   1,3,7-nonatriene, &szlig;-cedrene)) from the intercrops. If the   selected plant can have additional properties that meet other   farmer needs like increased nitrogen input in the soil or weed   suppression, these qualities should be promoted to achieve   multiple goals with only one plant. Such is the case of the   repellent plant <i><i>Desmodium </i></i>uncinatum, which has a series of   very astonishing properties. For example, the weed   suppressing property is achieved by a blend of secondary   metabolites in the root exudates that include seed germination   stimulants and at the same time post-germination inhibitors   resulting in &#8220;suicidal germination&#8221; (Tsanuo <i>et al</i>. 2003). Not   only its weed suppressive properties but also the fact that   <i><i>Desmodium </i></i>is a legume that increases nitrogen availability in   the soil that improves land productivity, and increases gross   cash returns (e.g. Khan <i>et al</i>. 2001) makes it highly attractive.   At the same time, farmers can use this species as a nutritious   and perennial fodder for cattle improving the productivity of   meat and milk. Moreover the seeds of <i>D. uncinatum</i> represent   a valuable commodity that has a local high demand among   different groups of farmers (Khan <i>et al</i>. 2000). Screening for   multiple properties can therefore increase the advantages of   diversifying a crop, by supplying natural fertilizers, herbicides,   pesticides and also providing fodder for cattle.</p>     <p>   But development of the push-pull strategy does not end   here. Khan <i>et al</i>. (2008a) also studied the adoption of this   practice as a technological package by farmers, showing that   by 2007 it was already adopted by thousands of farmers in   eastern Africa and the program is still expanding (Khan <i>et al</i>.   2008b). The implementation of this push-pull technology has   been shown to increase maize yields by 30%, providing the   best evidence that diversification practices are useful in   managing pests, increasing yield and moreover giving farmers   the possibility of additional income, without an intensive use   of pesticides.</p>     <p><b>   <font size="3">Summary</font></b></p>     <p>   Our literature review revealed contradictory effects of increased   diversity on natural enemies, herbivores and production, and   the expected results of reduced pest damage were only achieved   in 50% of the cases. However, some examples demonstrate   that diversification practices can translate into a successful   management technology that is adopted by thousands of   farmers. The current available data suggest a series of steps   that should be taken to design successful and competitive   diversification practices that can be adopted by the farmers:</p>     <p>   - Gather precise information on the natural history of the   pest and their natural enemies to selectively provide resources   and shelter for the natural enemies, but not for the pest.</p>     <p>   - Take into account the farmer&#8217;s needs to choose the   &#8220;right&#8221; plant(s).</p>     ]]></body>
<body><![CDATA[<p>   - Be open in the search for the appropriate functional plant   and screen as many plants as possible.</p>     <p>   - Favor plants that fulfill more than one function at the   same time.</p>     <p>   - Evaluate the effect of the chosen plant(s) on pests, natural   enemies, crop damage, crop development and yield.</p>     <p>   - Study the effectiveness of different arrangement patterns.</p>     <p>   - Perform comparative field experiments at different   locations and in different years to define the limitations of the   proposed practice.</p>     <p>   - Perform an economic study comparing the conventional   methods with the proposed practice.</p>     <p>   - Evaluate the labor intensity of the practice and the   willingness of the farmer to implement it.</p>     <p>   - Reach a mechanistic understanding of how the selected   plant achieves the expected results to reinforce those   characteristics on the selected plants or search for them in other   plants.</p>     <p>   - Test if the combination of several different functional   plants leads to a synergistic effect on pest suppression and crop   yield.</p>     <p>   - Distribute the knowledge among farmers, including onfarm   experiments where farmers evaluate and quantify the   effectiveness of the practice. </p>     ]]></body>
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