Ethylene rapidly up-regulates the activities of both monomeric GTP-binding proteins and protein kinase(s) in epicotyls of pea

Igor E. Moshkov, Galina V. Novikova, Luis A.J. Mur, Aileen R. Smith, and Michael A. Hall. (2003). Ethylene rapidly up-regulates the activities of both monomeric GTP-binding proteins and protein kinase(s) in epicotyls of pea. Plant Physiology, 131(4), 1718-1726 RAE2008

Cross-species identification of Mendel's/locus

Armstead, I. P., Donnison, I. S., Aubry, S., Harper, J. A., H?rtensteiner, S., James, C. L., Mani, J., Moffet, M., Ougham, H. J., Roberts, L. A., Thomas, A. M., Weeden, N., Thomas, S., King, I. P. (2007). Cross-species identification of Mendel's/locus. Science, 315 (5808), 73. Sponsorship: BBSRC RAE2008

Geography and Genetics of Ecological Speciation in Pea Aphids

During ecological speciation, divergent natural selection drives evolution of ecological specialization and genetic differentiation of populations on alternate environments. Populations diverging onto the same alternate environments may be geographically widespread, so that divergence may occur at an array of locations simultaneously. Spatial variation in the process of divergence may produce a pattern of differences in divergence among locations called the Geographic Mosaic of Divergence. Diverging populations may vary in their degree of genetic differentiation and ecological specialization among locations. My dissertation examines the pattern and evolutionary processes of divergence in pea aphids (Acyrthosiphon pisum) on alfalfa (Medicago sativa) and clover (Trifolium pretense). In Chapter One, I examined differences among North American aphid populations in genetic differentiation at nuclear, sequence-based markers and in ecological specialization, measured as aphid fecundity on each host plant. In the East, aphids showed high host-plant associated ecological specialization and high genetic differentiation. In the West, aphids from clover were genetically indistinguishable from aphids on alfalfa, and aphids from clover were less specialized. Thus, the pattern of divergence differed among locations, suggesting a Geographic Mosaic of Divergence. In Chapter Two, I examined genomic heterogeneity in divergence in aphids on alfalfa and clover across North America using amplified fragment length polymorphisms (AFLPs). The degree of genetic differentiation varied greatly among markers, suggesting that divergent natural selection drives aphid divergence in all geographic locations. Three of the same genetic markers were identified as evolving under divergent selection in the eastern and western regions, and additional divergent markers were identified in the East. In Chapter Three, I investigated population structure of aphids in North America, France, and Sweden using AFLPs. Aphids on the same host plant were genetically similar across many parts of their range, so the evolution of host plant specialization does not appear to have occurred independently in every location. While aphids on alfalfa and clover were genetically differentiated in most locations, aphids from alfalfa and clover were genetically similar in both western North America and Sweden. High gene flow from alfalfa onto clover may constrain divergence in these locations.

Calibración del modelo Cropsyst para simular el balance hídrico en cultivos invernales del centro oeste de Entre Ríos

El trabajo tiene por objetivo calibrar el modelo CropSyst para simular la producción de materia seca y el balance hídrico de los cultivos de trigo (Triticum aestivum L.) ciclo largo y ciclo corto, colza (Brassica napus), lino (Linum usitatissimum) y arveja proteica (Pisum sativum) en el centro oeste de Entre Ríos. Adicionalmente, se incluyó un análisis de sensibilidad de parámetros que podrían tener, a priori, un mayor impacto sobre el crecimiento y desarrollo y en especial sobre el balance hídrico. La optimización de Cropsyst se realizó utilizando datos obtenidos de un experimento a campo, realizado con un diseño en bloques completos al azar, con cuatro repeticiones. Los parámetros medidos fueron: fenología, humedad del suelo, radiación interceptada por el cultivo, biomasa, índice de área foliar, área foliar específica y rendimiento. Durante el proceso de calibración se utilizó un procedimiento iterativo para optimizar las salidas de crecimiento y desarrollo de los cultivos. Una vez optimizado el modelo agronómico se procedió a evaluar las predicciones relacionadas con el contenido de agua (cm-3 cm-3) por capas y la lámina de agua total (mm) hasta una profundidad de 50 cm. En relación con el contenido de agua se tomaron dos fechas aproximadas de observación y registro, seleccionadas en base al período crítico de cada especie y la lámina de agua total se evaluó durante todo el ciclo de los cultivos. A los efectos de simular la infiltración de agua en el suelo se usaron dos métodos incluidos en modelo: el método en cascada y el método de las diferencias finitas. En general, Cropsyst brindó estimaciones razonables en cuanto a la biomasa, intercepción de radiación y rendimiento. Las predicciones del contenido de agua en el suelo para cada profundidad resultaron poco satisfactorias. Sin embargo, cuando se consideró la lámina de agua a 50 cm. las predicciones fueron más aceptables. Con relación al análisis de sensibilidad, los parámetros evaluados no mostraron cambios significativos en la simulación del agua en el suelo ni en el resto de los indicadores analizados, sugiriendo que mejores ajustes en la predicción del contenido de agua deberían lograrse seleccionando otro tipo de parámetros. La revisión y adaptación de las funciones de pedo-transferencia para estimar los límites máximos y mínimos de almacenaje de agua en el suelo probablemente mejore los resultados obtenidos en este trabajo.

Interacción entre fecha y densidad de siembra en cultivos invernales

El cultivo de soja (Glycine max L.) de primera predomina en las secuencias agrícolas de la provincia de Entre Ríos, desperdiciándose una enorme cantidad de recursos al realizar un sólo cultivo por año. El estudio comparado de la interacción entre fecha de siembra y densidad en cultivos invernales ayudaría a identificar la mejor combinación para maximizar el rendimiento, alentando la intensificación de la secuencia de producción. El objetivo de este trabajo fue estudiar de manera comparada la interacción entre la fecha de siembra y la densidad de siembra en arveja proteica (Pisum sp.), lino (Linum usitatissimum L.), trigo ciclo corto y ciclo largo (Triticum aestivum L.). El experimento fue conducido en la EEA Paraná del INTA durante la campaña 2008/09, se evaluaron tres fechas de siembra (FS) y los tratamientos consistieron en una combinación factorial de cultivos invernales y densidades de siembra (DS). Se determinó la biomasa total acumulada hasta madurez, el rendimiento, los componentes y subcomponentes del rendimiento. Además, en la fecha de siembra más temprana se cuantificaron los componentes ecofisiólogicos condicionantes de la acumulación de biomasa, i.e. intercepción de la radiación fotosinteticamente activa (RFA), eficiencia en el uso de la radiación (EUR) y de partición. La eficiencia de intercepción de RFA como así también la EUR difirió significativamente en respuesta a la densidad de siembra, pero no fue significativo su efecto sobre la RFA acumulada y la evolución de la biomasa durante el ciclo de los cultivos. En ninguno de los cultivos estudiados existió interacción significativa FSxDS sobre el rendimiento o sus componentes numéricos, sólo en los cultivares de trigo se registraron efectos significativos de la interacción FSxDS sobre los subcomponentes del rendimiento. Para estudiar mejor la interacción FSxDS en especies con diferente plasticidad sería conveniente explorar su comportamiento sin limitaciones hídricas, complementando dichos estudios con la medición de variables ecofisológicas

Acción del cloropropham en la actividad argininadecarboxilasa de semillas de arveja en germinación

p.59-68

Arsenite transport into paddy rice (Oryza sativa) roots

Here the mechanism of arsenite transport into paddy rice (Oryza sativa) roots, uptake of which is described by Michaelis-Menten kinetics, is reported. A recent study on yeast (Saccharomyces cerevisiae) showed that undissociated arsenite (its pK_a is 9.2) was transported across the plasma membrane via a glycerol transporting channel. To investigate whether the same mechanism of transport was involved for rice, competitive studies with glycerol, which is transported into cells via aquaporins, were performed. Glycerol competed with arsenite for transport in a dose-dependent manner, indicating that arsenite and glycerol uptake mechanisms were the same. Arsenate transport was unaffected by glycerol, confirming that arsenate and arsenite are taken up into cells by different mechanisms. Antimonite, an arsenite analogue that is transported into S. cerevisiae cells by aquaporins, also competed with arsenite transport in a dose-dependent manner, providing further evidence that arsenite is transported into rice roots via glycerol transporting channels. Mercury (Hg²⁺) inhibited both arsenite and arsenate uptake, suggesting that inhibition of influx was due to general cellular stress rather than the specific action of Hg²⁺ on aquaporins. Arsenite uptake by pea (Pisum sativum) and wheat (Triticum aestivum) was also described by Michaelis-Menten kinetics.

Herbicidal activity of volatiles from coriander, winter savory, cotton lavender, and thyme isolated by hydrodistillation and supercritical fluid extraction

The volatiles from Coriandrum sativum L., Satureja montana L., Santolina chamaecyparissus L., and Thymus vulgaris L. were isolated by hydrodistillation (essential oil) and supercritical fluid extraction (volatile oil). Their effect on seed germination and root and shoot growth of the surviving seedlings of four crops (Zea mays L., Triticum durum L., Pisum sativum L., and Lactuca sativa L.) and two weeds (Portulaca oleracea L. and Vicia sativa L.) was investigated and compared with those of two synthetic herbicides, Agrocide and Prowl. The volatile oils of thyme and cotton lavender seemed to be promising alternatives to the synthetic herbicides because they were the least injurious to the crop species. The essential oil of winter savory, on the other hand, affected both crop and weeds and can be appropriate for uncultivated fields.

Studies on arbuscular mycorrhiza (AM) in the Alentejo (Portugal) using pea mutants resistant to AM fungi as a control tool for field conditions

The utilization and management of arbuscular mycorrhiza (AM) symbiosis may improve production and sustainability of the cropping system. For this purpose, native AM fungi (AMF) were sought and tested for their efficiency to increase plant growth by enhanced P uptake and by alleviation of drought stress. Pot experiments with safflower (Carthamus tinctorius) and pea (Pisum sativum) in five soils (mostly sandy loamy Luvisols) and field experiments with peas were carried out during three years at four different sites. Host plants were grown in heated soils inoculated with AMF or the respective heat sterilized inoculum. In the case of peas, mutants resistant to AMF colonization were used as non-mycorrhizal controls. The mycorrhizal impact on yields and its components, transpiration, and P and N uptake was studied in several experiments, partly under varying P and N levels and water supply. Screening of native AMF by most probable number bioassays was not very meaningful. Soil monoliths were placed in the open to simulate field conditions. Inoculation with a native AMF mix improved grain yield, shoot and leaf growth variables as compared to control. Exposed to drought, higher soil water depletion of mycorrhizal plants resulted in a haying-off effect. The growth response to this inoculum could not be significantly reproduced in a subsequent open air pot experiment at two levels of irrigation and P fertilization, however, safflower grew better at higher P and water supply by multiples. The water use efficiency concerning biomass was improved by the AMF inoculum in the two experiments. Transpiration rates were not significantly affected by AM but as a tendency were higher in non-mycorrhizal safflower. A fundamental methodological problem in mycorrhiza field research is providing an appropriate (negative) control for the experimental factor arbuscular mycorrhiza. Soil sterilization or fungicide treatment have undesirable side effects in field and greenhouse settings. Furthermore, artificial rooting, temperature and light conditions in pot experiments may interfere with the interpretation of mycorrhiza effects. Therefore, the myc- pea mutant P2 was tested as a non-mycorrhizal control in a bioassay to evaluate AMF under field conditions in comparison to the symbiotic isogenetic wild type of var. FRISSON as a new integrative approach. However, mutant P2 is also of nod- phenotype and therefore unable to fix N2. A 3-factorial experiment was carried out in a climate chamber at high NPK fertilization to examine the two isolines under non-symbiotic and symbiotic conditions. P2 achieved the same (or higher) biomass as wild type both under good and poor water supply. However, inoculation with the AMF Glomus manihot did not improve plant growth. Differences of grain and straw yields in field trials were large (up to 80 per cent) between those isogenetic pea lines mainly due to higher P uptake under P and water limited conditions. The lacking N2 fixation in mutants was compensated for by high mineral N supply as indicated by the high N status of the pea mutant plants. This finding was corroborated by the results of a major field experiment at three sites with two levels of N fertilization. The higher N rate did not affect grain or straw yields of the non-fixing mutants. Very efficient AMF were detected in a Ferric Luvisol on pasture land as revealed by yield levels of the evaluation crop and by functional vital staining of highly colonized roots. Generally, levels of grain yield were low, at between 40 and 980 kg ha-1. An additional pot trial was carried out to elucidate the strong mycorrhizal effect in the Ferric Luvisol. A triplication of the plant equivalent field P fertilization was necessary to compensate for the mycorrhizal benefit which was with five times higher grain yield very similar to that found in the field experiment. However, the yield differences between the two isolines were not always plausible as the evaluation variable because they were also found in (small) field test trials with apparently sufficient P and N supply and in a soil of almost no AMF potential. This similarly occurred for pea lines of var. SPARKLE and its non-fixing mycorrhizal (E135) and non-symbiotic (R25) isomutants, which were tested in order to exclude experimentally undesirable benefits by N2 fixation. In contrast to var. FRISSON, SPARKLE was not a suitable variety for Mediterranean field conditions. This raises suspicion putative genetic defects other than symbiotic ones may be effective under field conditions, which would conflict with the concept of an appropriate control. It was concluded that AMF resistant plants may help to overcome fundamental problems of present research on arbuscular mycorrhiza, but may create new ones.

Root effects on the turnover of grain legume residues in soil

Das Ziel dieser Arbeit war, die Einflüsse von Wurzeln und Rhizodeposition auf den Umsatz von Körnerleguminosenresiduen und damit verknüpfte mikrobielle Prozesse zu untersuchen. In einem integrierten Versuch wurden Ackerbohne (Vicia faba L.), Erbse (Pisum sativum L.) und Weiße Lupine (Lupinus albus L.) untersucht. Der Versuch bestand aus drei Teilen, zwei Gefäß-Experimenten und einem Inkubationsexperiment, in denen ausgehend von einem Gefäß-Experiment derselbe Boden und dasselbe Pflanzenmaterial verwendet wurden. In Experiment I wurde die Stickstoff-Rhizodeposition der Körnerleguminosenarten, definiert als wurzelbürtiger N nach dem Entfernen aller sichtbaren Wurzeln im Boden, gemessen und der Verbleib des Rhizodepositions-N in verschiednenen Bodenpools untersucht. Dazu wurden die Leguminosen in einem Gefäßversuch unter Verwendung einer in situ 15N-Docht-Methode mit einer 15N Harnstofflösung pulsmarkiert. In Experiment II wurde der Umsatz der N-Rhizodeposition der Körnerleguminosen und der Einfluss der Rhizodeposition auf den anschließenden C- und N-Umsatz der Körnerleguminosenresiduen in einem Inkubationsexperiment untersucht. In Experiment III wurde der N-Transfer aus den Körnerleguminosenresiduen einschließlich N-Rhizodeposition in die mikrobielle Biomasse und die Folgefrüchte Weizen (Triticum aestivum L.) und Raps (Brassica napus L.) in einem Gewächshaus-Gefäßversuch ermittelt. Die in situ 15N Docht-Markierungs-Methode wies hohe 15N Wiederfindungsraten von ungefähr 84 Prozent für alle drei Leguminosenarten auf und zeigte eine vergleichsweise homogene 15N Verteilung zwischen verschiedenen Pflanzenteilen zur Reife. Die Wurzeln zeigten deutliche Effekte auf die N-Dynamik nach dem Anbau von Körnerleguminosen. Die Effekte konnten auf die N-Rhizodeposition und deren anschließenden Umsatz, Einflüsse der Rhizodeposition von Körnerleguminosen auf den anschließenden Umsatz ihrer Residuen (Stängel, Blätter, erfassbare Wurzeln) und die Wirkungen nachfolgender Nichtleguminosen auf den Umsatzprozess der Residuen zurückgeführt werden: Die N-Rhizodeposition betrug zur Reife der Pflanzen bezogen auf die Gesamt-N- Aufnahme 13 Prozent bei Ackerbohne und Erbse und 16 Prozent bei Weißer Lupine. Bezogen auf den Residual N nach Ernte der Körner erhöhte sich der relative Anteil auf 35 - 44 Prozent. Die N-Rhizodeposition ist daher ein wesentlicher Pool für die N-Bilanz von Körnerleguminosen und trägt wesentlich zur Erklärung positiver Fruchtfolgeeffekte nach Körnerleguminosen bei. 7 - 21 Prozent des Rhizodepositions-N wurden als Feinwurzeln nach Nasssiebung (200 µm) wiedergefunden. Nur 14 - 18 Prozent des Rhizodepositions-N wurde in der mikrobiellen Biomasse und ein sehr kleiner Anteil von 3 - 7 Prozent in der mineralischen N Fraktion gefunden. 48 bis 72 Prozent der N-Rhizodeposition konnte in keinem der untersuchten Pools nachgewiesen werden. Dieser Teil dürfte als mikrobielle Residualmasse immobilisiert worden sein. Nach 168 Tagen Inkubation wurden 21 bis 27 Prozent des Rhizodepositions-N in den mineralisiert. Der mineralisierte N stammte im wesentlichen aus zwei Pools: Zwischen 30 Prozent und 55 Prozent wurde aus der mikrobiellen Residualmasse mineralisiert und eine kleinere Menge stammte aus der mikrobielle Biomasse. Der Einfluss der Rhizodeposition auf den Umsatz der Residuen war indifferent. Durch Rhizodeposition wurde die C Mineralisierung der Leguminosenresiduen nur in der Lupinenvariante erhöht, wobei der mikrobielle N und die Bildung von mikrobieller Residualmasse aus den Leguminosenresiduen in allen Varianten durch Rhizodepositionseinflüsse erhöht waren. Das Potential des residualen Körnerleguminosen-N für die N Ernährung von Folgefrüchten war gering. Nur 8 - 12 Prozent des residualen N wurden in den Folgenfrüchten Weizen und Raps wiedergefunden. Durch die Berücksichtigung des Rhizodepositions-N war der relative Anteil des Residual-N bezogen auf die Gesamt-N-Aufnahme der Folgefrucht hoch und betrug zwischen 18 und 46 Prozent. Dies lässt auf einen höheren N-Beitrag der Körnerleguminosen schließen als bisher angenommen wurde. Die residuale N-Aufnahme von Weizen von der Blüte bis zur Reife wurde durch den Residual-N gespeist, der zur Blüte in der mikrobiellen Biomasse immobilisiert worden war. Die gesamte Poolgröße, Residual-N in der mikrobiellen Biomasse und in Weizen, veränderte sich von der Blüte bis zur Reife nicht. Jedoch konnte ein Rest von 80 Prozent des Residual-N in keinem der untersuchten Pools nachgewiesen werden und dürfte als mikrobielle Residualmasse immobilisiert worden sein oder ist noch nicht abgebaut worden. Die zwei unterschiedlichen Folgefrüchte - Weizen und Raps - zeigten sehr ähnliche Muster bei der N-Aufnahme, der Residual-N Wiederfindung und bei mikrobiellen Parametern für die Residuen der drei Körnerleguminosenarten. Ein differenzierender Effekt auf den Umsatz der Residuen bzw. auf das Residual-N-Aneignungsvermögen der Folgefrüchte konnte nicht beobachtet werden.

Survey of Agricultural Practices and Alternatives to Pesticide Use to Conserve Water Resources in the Mojanda Watershed, Ecuador

Agriculture in the Mojanda Watershed is facing rainfall reductions caused by climate change. Reductions of water availability in the Watershed are also due to constant extension of the agricultural activities into the páramo ecosystem above 3000m a.s.l., with this ecosystem having immanently important functions in the local water balance. The application of pesticides threatens the quality of water and with less precipitation contaminations will further concentrate in the outflow. To analyze problems associated with agricultural practices in the area a questionnaire about agricultural practices (28) was conducted and fields (20) were surveyed for pests and diseases with a focus on potatoes (Solanum tuberosum L.), tree tomatoes (Solanum betaceum Cav.) and peas (Pisum sativum L.). Potatoes were infected to a low degree with Phytophthora infestans and according to the farmers the Andean potato weevil (Premnotrypes spec.) caused biggest losses. To combat the weevil the soils are disinfected with toxic Carbofuran (WHO Class 1B). Tree tomatoes showed symptoms of various fungal diseases. Most important was Fusarium solani causing the branches to rot and Anthracnosis (Colletotrichum gloeosporioides) causing the fruits to rot. Fungicide applications were correspondingly high. Peas were only minorly affected by Ascochyta blight (Mycosphaerella pinodes) and a root rot. Overall 19 active ingredients were applied of which fungicide Mancozeb (WHO class table 5) and insecticide Carbofuran (WHO Class 1B) were applied the most. Approved IPM methods are advised to reduce pesticide use. For tree tomatoes regular cutting of branches infected with F. solani and regular collection and disposal of infected fruits with Anthracnosis are advised. For potatoes plastic barriers around the fields prevent the Andean potato weevil from laying eggs thus reducing infestation with the larvae in the tubers. Local bioinsecticide “Biol” seems effective and without harm to the environment, although not used by many farmers. Organic fertilization promises to restore decreasing soil fertility, water holding capacity and reduce erosion. The here presented alternatives and strategies to reduce pesticide use pose an opportunity to preserve the water resources of the region.

Pea-barley intercrops use nitrogen sources 20-30% more efficiently than the sole crops

Field pea (Pisum sativum L.) and spring barley (Hordeum vulgare L) were intercropped and sole cropped to compare the effects of crop diversity on the use of nitrogen sources in European organic crop-ping systems. Across a wide range of growing condi-tions pea-barley intercropping showed that nitrogen sources were used from 17 to 31% more efficiently by the intercrop than by the sole crops. Intercropping technologies offers the opportunity for organic cropping systems to utilize N complementarity between component crops, without compromising total crop N yield levels

Mutant alleles at the rugosus loci in pea affect seed moisture sorption isotherms and the relations between seed longevity and moisture content

Pea (Pisum sativum L.) mutant near-isogenic lines (RRrbrb, rrRbRb, rrrbrb) with lower starch but higher lipid contents, brought about by lesions in the starch biosynthetic pathway, had seed moisture sorption isotherms displaced below that of the wild type (RRRbRb). The negative logarithmic relationship between seed longevity and seed storage moisture content (%, f.wt basis), determined in hermetic storage at 65 degreesC, also differed: longevity in the mutant near-isogenic lines was poorer and less sensitive to moisture content than in the wild type (i.e. C-w was lower). The low-moisture-content limit (m(c)) to this relation also differed, being lower in the mutant near-isogenic lines (5.4-5.9%) than in the wild type (6.1%). In contrast, all four near-isogenic lines showed no difference (P >0.25) in the negative semilogarithmic relationship between equilibrium relative humidity (ERH) and seed longevity. It is concluded that the effect of these alleles at the r and rb loci on seed longevity. was largely indirect; a consequence of their effect on seed composition and hence on moisture sorption isotherms. However, this explanation could not be invoked at moisture contents below mc where differences in longevity remained substantial (RRRbRb double that of rrrbrb). Hence, these mutant alleles affected seed longevity directly at very low moisture contents.

Intraspecific heritable variation in life-history traits can alter the outcome of interspecific competition among insect herbivores

Competition is one of the most important biotic factors determining the structure of ecological communities. In this study, we show that there is variation in competitive ability between two clones of the pea aphid, Acyrthosiphon pisum, both of which out-compete a clone of the vetch aphid, Megoura viciae, in the laboratory. We tested whether this variation in competitive ability would alter the outcome of interspecific competition in the field. White one pea aphid clone followed the pattern set in the laboratory, out-competing the Megoura viciae clone, another showed the reverse effect with Megoura viciae dominating. These differences appear to be the result of variation in early population growth rate between the pea aphid clones, rather than predation, although predation did lead to the eventual extinction of colonies. We also questioned whether intra- and interspecific differences in predator escape behaviour could affect the outcome of competition in the field. All three clones responded similarly to the presence of foraging hoverfly larvae (Episyrphus balteatus), but the Megoura viciae clone dropped from the plant significantly less often in response to the presence of a foraging two-spot ladybird (Adalia bipunctata). This work provides evidence that intraspecific variation in competitive ability can alter the outcome of interspecific competitive interactions in nature and suggests that species-specific behavioural. traits may have the potential to modify the outcome of these interactions. (c) 2005 Gesellschaft fur Okologie. Published by Elsevier GmbH. All rights reserved.

Resistance is costly: trade-offs between immunity, fecundity and survival in the pea aphid

Parasitoids are among the most important natural enemies of insects in many environments. Acyrthosiphon pisum, the pea aphid, is a common pest of the leguminous crops in temperate regions. Pea aphids are frequently attacked by a range of endoparasitic wasps, including the common aphidiine, Aphidius ervi. Immunity to parasitoid attack is thought to involve secondary symbiotic bacteria, the presence of which is associated with the death of the parasitoid egg. It has been suggested that there is a fecundity cost of resistance, as individuals carrying the secondary symbionts associated with parasitoid resistance have fewer offspring. Supporting this hypothesis, we find a positive relationship between fecundity and susceptibility to parasitoid attack. There is also a negative relationship between fecundity and off-plant survival time (which positively correlates with resistance to parasitoid attack). Taken together, these results suggest that the aphids can either invest in defence (parasitoid resistance, increased off-plant survival time) or reproduction, and speculate that this may be mediated by changes in the aphids' endosymbiont fauna. Furthermore, there is a positive relationship between aphid size and resistance, suggesting that successful resistance to parasitoid attack may involve physical, as well as physiological, defences.