Lepidopteran pest populations and crop yields in row intercropped broccoli

Abstract

• 1 To determine how the presence of non‐host plants impact populations of lepidopteran pests and yield of broccoli, Brassica oleracea var. italica L., broccoli was intercropped with tomato, Lycopersicon esculentum Miller, or yellow sweetclover (YSC), Melilotus officinalis L., and compared with broccoli monoculture grown at two densities (high and low).
• 2 During a 1997 study, numbers of early instar imported cabbageworm, Artogeia rapae L., were greater in broccoli monoculture compared with intercropped plantings. In contrast, late instar A. rapae numbers were significantly higher in intercropped broccoli during most of the season, indicating greater survival of early instar stages in intercropped habitats.
• 3 Cabbage looper, Trichoplusia ni Hübner, eggs and larvae were more abundant in broccoli monoculture than intercropped broccoli in 1997. In addition, T. ni populations were found at significantly lower levels in broccoli‐YSC compared with broccoli‐tomato habitat throughout the season.
• 4 During a 1998 study, numbers of early and late A. rapae instars were greater in intercropped broccoli on most sampling dates.
• 5 The mean percentages of harvested broccoli heads infested with insects and associated frass were higher in monocultures than intercropped treatments in 1997. Additionally, a significantly lower number of contaminants were found in the crowns of broccoli harvested from broccoli‐YSC (YSC) compared with broccoli‐tomato (tomato) habitat. However, broccoli heads were significantly smaller in intercropped habitats.
• 6 Variations in insect responses and challenges to using crop diversification as a management tool are discussed.

     

Population densities of herbivorous lepidopterans in diverse cruciferous cropping habitats: Effects of mixed cropping and using a living mulch

Two field experiments were conducted in Hawaii to determine the responses of lepidopteran pests and their associated natural enemies to broccoli (Brassica oleracea), grown in monoculture or interplanted with chili pepper (Capsicum annuum), or yellow sweetclover (YSC) (Melilotus officinalis). The population responses of Artogeia rapae and Trichoplusia ni to habitat types were not consistent during the field trials. Additionally, their reactions to habitat diversification differed according to non-host plant species. For example, Trichoplusia ni densities were significantly lower in the broccoli–YSC habitat contrasted with the broccoli–pepper habitat on most sampling dates. Similarly, the proportion of plants damaged by Hellula undalis was significantly lower in the broccoli–YSC habitat contrasted with broccoli–pepper habitat. Findings from the study, suggest that biological control agents (e.g., spiders, parasitoids, and␣entomopathogens) are the most significant factors influencing the abundance of A.␣rapae in habitat type and that these agents inflict greater mortality on A. rapae in diculture than monoculture habitats.     

Effects of crop diversification levels and fertilization regimes on abundance of Brevicoryne brassicae (L.) and its parasitization by Diaeretiella rapae (M’Intosh) in broccoli

1 The effects of intercropping via competition on crop yields, pest [cabbage aphid Brevicoryne brassicae (L.)] abundance, and natural enemy efficacy were studied in the Brassica oleracea L. var. italica system. 2 From May to December 2004, insect populations and yield parameters were monitored in summer and autumn in broccoli monoculture and polyculture systems with or without competition from Brassica spp. (mustard), or Fagopyrum esculentum Moench (buckwheat), with addition of organic (compost) or synthetic fertilizer. 3 Competition from buckwheat and mustard intercrops did not influence pest density on broccoli; rather, aphid pressure decreased and natural enemies of cabbage aphid were enhanced in intercropping treatments, but this varied with the intercropped plant and season (summer vs. autumn). 4 In compost‐fertilized broccoli systems, seasonal parasitization rates of B. brassicae by Diaeretiella rapae (M’Intosh) increased along with the expected lower aphid pressure compared with synthetically fertilized plants.     

Neighbouring monocultures enhance the effect of intercropping on the turnip root fly (Delia floralis)

Knowledge of insect behaviour is essential for accurately interpreting studies of diversification and to develop diversified agroecosystems that have a reliable pest‐suppressive effect. In this study, we investigated the egg‐laying behaviour of the turnip root fly, Delia floralis (Fall.) (Diptera: Anthomyiidae), in an intercrop‐monoculture system. We examined both the main effect of intercropping and the effect on oviposition in the border zone between a cabbage monoculture [Brassica oleracea L. var. capitata (Brassicaceae)] and a cabbage‐red clover intercropping system [Trifolium pratense L. (Fabaceae)]. To investigate the border‐effect, oviposition was measured along a transect from the border between the treatments to the centre of experimental plots. Intercropping reduced the total egg‐laying of D. floralis with 42% in 2003 and 55% in 2004. In 2004, it was also found that the spatial distribution of eggs within the experimental plots was affected by distance from the adjoining treatment. The difference in egg‐laying between monoculture and intercropping was most pronounced close to the border, where egg‐laying was 68% lower on intercropped plants. This difference in egg numbers decreased gradually up to a distance of 3.5 m from the border, where intercropped plants had 43% fewer eggs than the corresponding monocultured plants. The reason behind this oviposition pattern is most likely that flies in intercropped plots have a higher probability of entering the monoculture if they are close to the border than if they are in the centre of a plot. When entering the monoculture, flies can pursue their egg‐laying behaviour without being disrupted by the clover. As the final decision to land is visually stimulated, flies could also be attracted to fly from the intercropped plots into the monoculture, where host plants are more visually apparent. Visual cues could also hinder flies in a monoculture from entering an intercropped plot. Other possible patterns of insect attack due to differences in insect behaviour are discussed, as well as the practical application of the results of this study.     

Seasonal differences in mineral content, distribution and leakage of sweet pepper seeds

Pepper seed quality is sensitive to variations in climatic conditions during seed development, which might be associated with accumulation, distribution and leakage of mineral elements from the seeds. This was examined in hybrid seeds of sweet pepper (Capsicum annuum L. cv.‘Hazera’ 1195) in two experiments during two growing seasons. The mean daily temperature (day/night) and daily total radiation receipt during seed development were 27.9/23.2°C and 8.63 kW m^?2 in the summer and 18.3/ 14.9°C and 3.18 kW m^?2 in the winter, respectively. Seeds developed in the summer had lower percentage of seedling emergence and leaked a larger portion (45%) of their K content into the water medium than in winter‐developed seeds. Summer seeds accumulated more K and Cl, but less P, Mg, Ca and the weight ratio of linoleic acid to oleic acid was lower than in the winter seeds. The season did not significantly affect N, S and total fatty acids. The most abundant element on the seed coat surface was K in the summer and Ca in the winter seeds. The cotyledon and endosperm of the summer seeds contained relatively higher ratios of K and Ca and lower ratios of P and Mg than the winter seeds. Transportation of mineral nutrients appeared to be involved in the effect of heat and moisture stresses on emergence quality of the pepper seeds.     

Within‐field spatial distribution patterns of corn planthopper,Peregrinus maidis,and severity of hopperburn and Maize mosaic virus symptoms as influenced by sunn hemp intercropping

Habitat management (e.g., intercropping) may alter within‐field spatial distribution patterns of herbivores, from a typical pattern as observed in a monoculture, and may influence patterns of crop injury. Field trials were conducted to study the effect of intercropping maize, Zea mays L. (Poaceae), with sunn hemp, Crotalaria juncea L. (Fabaceae) strips on within‐field spatial distribution patterns of corn planthopper, Peregrinus maidis (Ashmead) (Hemiptera: Delphacidae), and combined severity of hopperburn and Maize mosaic virus (MMV) (Rhabdoviridae: Nucleorhabdovirus) symptoms. In each field trial, spatially explicit data on P. maidis counts and ratings of severity of symptoms were obtained by sampling maize plants at weekly intervals. These data were used to examine the spatial patterns of P. maidis and severity of symptoms in maize‐intercropped and monoculture plots with Spatial Analysis for Distance IndicEs (SADIE) methodology. Spatial aggregation patterns of P. maidis in each treatment plot were not consistent among the field trials and tended to be mediated by their population densities. Interpolation of local cluster indices showed that P. maidis were more often aggregated at the field edges, irrespective of treatment. At times of MMV incidence in field trials (fall 2010 and spring 2011), the patch clusters of P. maidis and symptomatic plants were located at the field edges, but were spatially unassociated in both treatment plots. The results provided an approximation of the unpredictability of P. maidis spatial patterns at different population densities and their association with severity of symptoms in two maize‐cropping systems. However, the gap clusters of symptomatic plants were primarily located at the field interiors and were larger in intercropped than in monoculture plots. Such spatial pattern of symptomatic plants resulted in the reduced incidence of MMV in the intercropped plot compared with the monoculture plot, suggesting intercropping sunn hemp can be a useful tool in the management of MMV in maize fields.     

Seed production and predispersal seed predation in the biennial composite species,Arctium minus (Hill) Bernh. and A. lappa L.

Summary Comparison was made of the seed production of two species of burdock, Arctium minus (Hill) Bernh. and A. lappa L., which are found in a variety of recently disturbed habitats throughout Waterloo County, Ontario. The mean number of seeds per plant of A. minus was significantly greater, 13,400 compared with 8,200 seeds in A. lappa. The mean number of heads per plant was also greater for A. minus than for A. lappa, 338 versus 112 heads respectively. Above-ground biomass allocated to reproductive heads was similar in A. minus (44.2%) and A. lappa (43.7%). Significant variation in the mean number of seeds per head was observed in A. minus, ranging from 29.0 to 47.1. The between-plant variation in this character is significantly greater than the variation between populations from various sites.Predispersal seed predation by the microlepidopteran Metzneria lappella (Fam. Gelechiidae) resulted in 28% to 71% loss of seeds of A. minus at different sites. The observed loss in A. lappa was dramatically lower at 15.5%. The number of seeds attacked by one larval predator varied from 2 to 15 with means from 7 to 9. In addition, great variation in the number of seeds and the number of predators per head was observed. Larvae were randomly distributed among the heads at 4 of 7 sites and aggregated at the other sites. Seed predation per head greater than 60% was found with the aggregated larval distributions. Only rarely was the number of seeds attacked by a larva associated with the number of larvae in the head. The winter dispersal of heads of A. minus was much lower than that for A. lappa, 62.2% and 21.0% respectively. Also, the dispersal of seeds from heads differed between the two species. Only 21.8% of the seeds of A. minus were lost compared with a seed dispersal rate of 56.5% for A. lappa.     

Inter‐seasonal persistence and size‐structuring of two minnow species within headwater streams in the Eastern Cape,South Africa

This study examined temporal variation in population dynamics and size structuring of two cyprinid minnows, Pseudobarbus afer and Barbus anoplus, in relation to their proximate physical habitats. Population estimates were determined using three‐pass depletion sampling during both summer and winter. The habitats were characterised by seasonal variation in all physico‐chemical conditions and spatial variation in substrata compositions. Whereas significant differences in population size were noted between seasons for B. anoplus, no differences were found between seasons for density and capture probability for either species. An increase in boulders was associated with increase in population size and density for P. afer; for B. anoplus, increased percentages of bedrock and bank vegetation were associated with an increase in population size and probability of capture, respectively. According to Canonical Correspondence Analysis, size structuring in P. afer was explained predominantly by seasonality, with smaller length classes associated with the seasonal variable of summer, while larger length classes were associated with pH that was higher in winter. By comparison, for B. anoplus, the habitat variables – bank vegetation and bedrock – accounted for much of the explained variance for size structuring. Recruitment appeared to be the major driver of size structuring for the two species; refugia, especially boulders and bank vegetation, also appeared to be important. Overall, the two species were adapted to the headwater streams that were generally variable in environmental conditions. Potential invasions by non‐native invasive fishes that occur within the mainstream habitats threaten these two species. Efforts should continue to protect these minnows from such invasions by constructing barriers to upstream migration of non‐native fishes into these headwater habitats.     

Impact of seasonality,storage of semen,and sperm head‐shape on whole tissue methylation and expression of methylation responsive candidate genes in swine placenta and fetal livers from summer and winter breedings

Epigenetics includes the study of external factors that can influence the expression of genes by altering the accessibility of DNA through methylation. To investigate the epigenetic influence of season, sperm head shape, and semen storage on placental and fetal tissues, pregnancies were generated in the summer or winter using boar semen from either least or most sperm head shape change, collected during cool or warm seasons, and stored as cooled‐extended or cryopreserved. The lowest (p < 0.05) ratios of 5‐methylcytosine to 5‐hydroxymethylcytosine activity (5mC:5hmC) in fetal liver were from summer breedings and in placental tissues from winter breedings. The relative expression of placental CDH1 tended ( p < 0.10) to be greater in placenta generated from cryopreserved semen or semen collected during cool periods. The relative expression of placental GNAS was affected ( p < 0.05) by the interaction of breeding and semen collection seasons. Cryopreserved semen increased ( p < 0.05) the placental relative expression of GNAS. Placental MEST and RHOBTB3 tended ( p < 0.10) to have a greater relative expression from pregnancies generated using semen collected during cool periods used during winter breedings. Within fetal liver, the relative expression of GNAS and HGF was greater ( p < 0.05) from winter breedings. Interaction of winter breedings and least sperm head shape change tended ( p < 0.10) to have the greatest fetal liver expression of CDH1. Seasonality of semen collection, breeding, and the effect on sperm head shape change had an influence on the expression of genes with known differentially methylated regions or response to methylation activity from embryonic and extraembryonic tissues.     

Comparative effect of live food and manured treatments on water quality and production of ornamental carp, Cyprinus carpio var. koi L., during winter, summer, monsoon and post-monsoon growout experiments in concrete tanks

Four growth trials (each of 11 weeks duration) were conducted during different seasons throughout 1 year, namely, winter, summer, monsoon and post‐monsoon to assess the seasonal influence on growth and production of koi carp, Cyprinus carpio var. koi L. larvae in concrete tanks maintained under different management regimes. Individual weight gain, survival rate, fish deformities and number of marketable fish in each season were compared among four management regimes: (i) carp larvae fed with live zooplankton (LF); (ii) direct fertilization with poultry manure (PM); (iii) direct fertilization with cow dung (CD); and (iv) a control system (C), where a commercial diet containing 32% crude protein was applied. There were three replicates for each treatment. The LF treatment produced significantly higher weight gain, survival rate and number of marketable fish in all seasons (P < 0.05), compared with the other treatments, through maintenance of better water quality and greater abundance of zooplankton in the system. Fish deformities were highest in the C treatment in every season (P < 0.05). Water temperatures averaged 18.6°C (winter), 29.7°C (summer), 28.3°C (monsoon), and 26.5°C in the post‐monsoon trial. Average weight gain of koi carp achieved during the winter trial (2.01 g in C to 4.44 g in LF) was considerably lower than that achieved in the summer, monsoon or post‐monsoon trials. Survival rates of carp ranged from 70.5% (C) to 95.5% (LF) in the winter trial, and were considerably lower than the other seasons. The number of marketable fish was also lowest during the winter trial.     

Effect of larval brain extracts derived from three swallowtail butterflies,Papilio helenus L., P. machaon L. and P. memnon L., on seasonal morph development of P. xuthus L. (Lepidoptera: Papilionidae)

Adults of the three papilionid butterflies, Papilio helenus L., Papilio machaon L. and Papilio memnon L., exhibit seasonal diphenism comprising spring and summer morphs. To elucidate the physiological mechanism underlying seasonal morph development in papilionid butterflies, we investigated whether a cerebral factor showing summer‐morph‐producing hormone (SMPH) activity is present in the brain of three Papilio species using an assay system with chilled male short‐day pupae of P. xuthus L. When 2% NaCl extracts derived from 20 larval brains of the three species were injected into abdomens of chilled male short‐day pupae of P. xuthus, all recipients destined to develop into spring‐morph adults developed into summer‐ and intermediate‐morph adults. On the other hand, all recipients injected with distilled water as a control developed into spring‐morph adults. These results indicate that a cerebral factor showing SMPH activity is present in the larval brain of the three Papilio species. Additionally, all recipients injected with 2% NaCl extracts derived from 20 adult brains of Bombyx mori L. also developed into summer‐ and intermediate‐morph adults. The results revealed that SMPH or a cerebral factor showing SMPH activity is widely distributed among lepidopteran insects.     

Seasonal variation of the capture and field distribution of sterile Anastrepha ludens in Tamaulipas,Mexico

Anastrepha ludens (Loew) (Diptera: Tephritidae) is one of the most important citrus pest in Mexico. The sterile insect technique (SIT) is used against pest populations of fruit flies for suppression, eradication, containment and prevention to reduce damages in fruit‐growing areas. In this study, we analyzed the seasonal variation of captures and field distribution of sterile A. ludens released in different seasons of the year in north‐eastern Mexico. Chilled releases were conducted by air at constant densities per ha on a citrus area for a period of 32 weeks that included the coldest and warmest seasons that is winter, spring and summer. Multilure traps baited with torula yeast pellets were used to capture sterile flies. Fly capture data were compared over the three seasons and correlated with climate. The lowest number of captures of the sterile insect occurred in the summer and the highest in winter and spring. High and low temperatures were negatively correlated with fly captures. Field distribution was also negatively correlated with high temperatures in summer, but no relationships were observed in winter and spring. No relationships were observed between rainfall with capture and field distribution of sterile flies. These results indicate that summer is a season involving agro‐ecological and environmental constraints for the capture and field distribution of sterile flies. This study may be useful for enhancing release strategies and optimizing economic resources in north‐eastern Mexico. Further research on the behaviour of sterile flies under stressful environments is suggested.     

Sarcosaprophagous Diptera assemblages in natural habitats in central Spain: spatial and seasonal changes in composition

The composition and spatial distribution of sarcosaprophagous Diptera assemblages were studied using carrion‐baited traps along a bioclimatic gradient of natural habitats in central Spain throughout the different seasons during 1 year. Calliphoridae and Muscidae were the most abundant families, accounting for, respectively, 41.9% and 35.1% of all Diptera specimens collected. Other abundant families were Heleomyzidae (8.4%), Sarcophagidae (6.9%) and Piophilidae (5.1%). Fly assemblage compositions differed among bioclimatic levels, with Chrysomya albiceps (Wiedemann) (Diptera: Calliphoridae) being the dominant species in mesomediterranean habitats, Muscina levida (Harris) (Diptera: Muscidae) the dominant species in supramediterranean habitats, and Prochyliza nigrimana (Meigen) (Diptera: Piophilidae) the dominant species in oromediterranean habitats. Differences in assemblage composition were also found among seasons. Thermophobic species such as Calliphora vicina Robineau‐Desvoidy (Diptera: Calliphoridae) and some species of Heleomyzidae were well represented during autumn, winter and spring in the three bioclimatic levels sampled. By contrast, thermophilic species such as Ch. albiceps and Lucilia sericata (Meigen) (Diptera: Calliphoridae) and most Muscidae and Sarcophagidae species were more abundant during summer and in mesomediterranean habitats located at lower elevations. Knowledge of the preferences of some species for certain habitats may be of ecological and forensic value and may establish a starting point for further research.     

Impacts of multiple extreme winter warming events on sub‐Arctic heathland: phenology,reproduction, growth,and CO2 flux responses

Extreme weather events can have strong negative impacts on species survival and community structure when surpassing lethal thresholds. Extreme, short‐lived, winter warming events in the Arctic rapidly melt snow and expose ecosystems to unseasonably warm air (for instance, 2–10 °C for 2–14 days) but upon return to normal winter climate exposes the ecosystem to much colder temperatures due to the loss of insulating snow. Single events have been shown to reduce plant reproduction and increase shoot mortality, but impacts of multiple events are little understood as are the broader impacts on community structure, growth, carbon balance, and nutrient cycling. To address these issues, we simulated week‐long extreme winter warming events – using infrared heating lamps and soil warming cables – for 3 consecutive years in a sub‐Arctic heathland dominated by the dwarf shrubs Empetrum hermaphroditum, Vaccinium vitis‐idaea (both evergreen) and Vaccinium myrtillus (deciduous). During the growing seasons after the second and third winter event, spring bud burst was delayed by up to a week for E. hermaphroditum and V. myrtillus, and berry production reduced by 11–75% and 52–95% for E. hermaphroditum and V. myrtillus, respectively. Greater shoot mortality occurred in E. hermaphroditum (up to 52%), V. vitis‐idaea (51%), and V. myrtillus (80%). Root growth was reduced by more than 25% but soil nutrient availability remained unaffected. Gross primary productivity was reduced by more than 50% in the summer following the third simulation. Overall, the extent of damage was considerable, and critically plant responses were opposite in direction to the increased growth seen in long‐term summer warming simulations and the ‘greening’ seen for some arctic regions. Given the Arctic is warming more in winter than summer, and extreme events are predicted to become more frequent, this generates large uncertainty in our current understanding of arctic ecosystem responses to climate change.     

Biochemical characterization of carotenoids in two species of <Emphasis Type="Italic">Trentepohlia</Emphasis> (Trentepohliales,Chlorophyta)

Two species of Trentepohlia, i.e., Trentepohlia aurea and Trentepohlia cucullata were collected from walls and tree bark, respectively, at two different seasons in a year. The total carotenoid content in both the species is very high during winter but decreases significantly during summer. By spectroscopic analysis, it was found that. T. aurea and T. cucullata growing in natural habitats are rich sources of carotenoids. The individual carotenoids were separated, identified, and estimated by HPLC, and identified as β-carotene along with some other carotenoids, i.e., neoxanthin, lutein, β-cryptoxanthin, β,γ-carotene, β,ε-carotene (absent during summer).     

Oviposition behavior of orange wheat blossom midge on low- vs. high-ranked grass seed heads

The discovery of Sm1, a highly effective resistance (R) gene that targets the first instar of the orange wheat blossom midge, Sitodiplosis mosellana (Géhin) (Diptera: Cecidomyiidae), has created concerns about wheat midge adaptation. Strategies for delaying adaptation to Sm1 include the simultaneous deployment of a resistance trait targeting a different life stage, i.e., the ovipositing adult female. Previous studies have shown that adult females distinguish between wheat genotypes and seed head developmental stages and are attracted by volatiles from young wheat heads. We focused on what happens after the female lands on the seed head, comparing in three tests a seed head of the high‐ranked pre‐anthesis ‘Roblin’ wheat, Triticum aestivum L. (Poaceae), and a head of one of three lower‐ranked types: post‐anthesis ‘Roblin’, pre‐anthesis ‘Key’ wheat (T. aestivum), and pre‐anthesis ‘Robust’ barley (Hordeum vulgare L.). Within each test, high‐ and low‐ranked heads were presented in choice and no‐choice assays, with the behavior of wheat midge females scored every 5 min from 20:30 to 23:00 hours, under mid‐summer natural light conditions and sunset occurring between 20:50 and 21:20 hours. Head type influenced both proportions of females observed on the head and proportions of females probing with the ovipositor. Head*assay interactions occurred only in the test comparing wheat to barley, with barley reducing females observed on the wheat head and wheat increasing females probing on barley. Results indicate that the wheat midge female detects plant cues while examining the seed head and that this detection contributes to differences in egg counts.     

Field overwintering biology of Spiroplasma kunkelii (Mycoplasmatales: Spiroplasmataceae) and its vector Dalbulus maidis (Hemiptera: Cicadellidae)

We studied the corn stunt spiroplasma (CSS), Spiroplasma kunkelii (Mycoplasmatales: Spiroplasmataceae) and its vector the corn leafhopper Dalbulus maidis (Hemiptera: Cicadellidae) under field conditions in Mexico. We surveyed for the presence of CSS in D. maidis by using PCR on samples of adults collected during the 2000–01 and 2003–04 winter (dry) seasons from irrigated low‐elevation sites and un‐irrigated high‐elevation sites. Also, we determined the body size and number of mature eggs of D. maidis females collected during the dry season in 2004 and in females collected on maize seedlings in the first months (June and July) of the wet (summer) season in 2005. Our PCR results showed that CSS was present in leafhopper adults collected during the 2000–01 and 2003–04 dry seasons in irrigated low‐elevation sites. However, in un‐irrigated high‐elevation sites, CSS was present in corn leafhopper adults caught before, but not during, the dry seasons. In these un‐irrigated high‐elevation sites, adult leafhoppers without CSS were recovered during the first 2 months (November and December) of the dry season from the foliage of wild perennial grasses. Females collected on wild perennial grasses in December 2004 showed similar head width and wing length to females caught on maize seedlings in June 2005. However, females collected on maize seedlings in July 2005 had the widest heads, longest wings and highest number of mature eggs. The largest body size of these females that arrived in July 2005 indicates that they are immigrants and support the Roff’s hypothesis that insect migrants should be larger than nonmigrants.     

Leaf anatomical and photosynthetic acclimation to cool temperature and high light in two winter versus two summer annuals

Acclimation of foliar features to cool temperature and high light was characterized in winter (Spinacia oleracea L. cv. Giant Nobel; Arabidopsis thaliana (L.) Heynhold Col‐0 and ecotypes from Sweden and Italy) versus summer (Helianthus annuus L. cv. Soraya; Cucurbita pepo L. cv. Italian Zucchini Romanesco) annuals. Significant relationships existed among leaf dry mass per area, photosynthesis, leaf thickness and palisade mesophyll thickness. While the acclimatory response of the summer annuals to cool temperature and/or high light levels was limited, the winter annuals increased the number of palisade cell layers, ranging from two layers under moderate light and warm temperature to between four and five layers under cool temperature and high light. A significant relationship was also found between palisade tissue thickness and either cross‐sectional area or number of phloem cells (each normalized by vein density) in minor veins among all four species and growth regimes. The two winter annuals, but not the summer annuals, thus exhibited acclimatory adjustments of minor vein phloem to cool temperature and/or high light, with more numerous and larger phloem cells and a higher maximal photosynthesis rate. The upregulation of photosynthesis in winter annuals in response to low growth temperature may thus depend on not only (1) a greater volume of photosynthesizing palisade tissue but also (2) leaf veins containing additional phloem cells and presumably capable of exporting a greater volume of sugars from the leaves to the rest of the plant.     

Row Ratios of Intercropping Maize and Soybean Can Affect Agronomic Efficiency of the System and Subsequent Wheat

Intercropping is regarded as an important agricultural practice to improve crop production and environmental quality in the regions with intensive agricultural production, e.g., northern China. To optimize agronomic advantage of maize (Zea mays L.) and soybean (Glycine max L.) intercropping system compared to monoculture of maize, two sequential experiments were conducted. Experiment 1 was to screening the optimal cropping system in summer that had the highest yields and economic benefits, and Experiment 2 was to identify the optimum row ratio of the intercrops selected from Experiment 1. Results of Experiment 1 showed that maize intercropping with soybean (maize || soybean) was the optimal cropping system in summer. Compared to conventional monoculture of maize, maize || soybean had significant advantage in yield, economy, land utilization ratio and reducing soil nitrate nitrogen (N) accumulation, as well as better residual effect on the subsequent wheat (Triticum aestivum L.) crop. Experiment 2 showed that intercropping systems reduced use of N fertilizer per unit land area and increased relative biomass of intercropped maize, due to promoted photosynthetic efficiency of border rows and N utilization during symbiotic period. Intercropping advantage began to emerge at tasseling stage after N topdressing for maize. Among all treatments with different row ratios, alternating four maize rows with six soybean rows (4M:6S) had the largest land equivalent ratio (1.30), total N accumulation in crops (258 kg ha^-1), and economic benefit (3,408 USD ha^-1). Compared to maize monoculture, 4M:6S had significantly lower nitrate-N accumulation in soil both after harvest of maize and after harvest of the subsequent wheat, but it did not decrease yield of wheat. The most important advantage of 4M:6S was to increase biomass of intercropped maize and soybean, which further led to the increase of total N accumulation by crops as well as economic benefit. In conclusion, alternating four maize rows with six soybean rows was the optimum row ratio in maize || soybean system, though this needs to be further confirmed by pluri-annual trials.     

Biomass yield and nitrogen fixation of legumes monocropped and intercropped with rye and rotation effects on a subsequent maize crop

The research is focused on an ecologically sound and highly productive cultivation system for fodder and/or biomass for thermal power generation on the basis of winter legumes and maize as subsequent summer crop, managed without additional nitrogen fertiliser. Therefore the yield of biomass and N-fixing capacity of a winter pea (Pisum sativum L.) and crimson clover (Trifolium incarnatum L.) monocropped and intercropped with rye (Secale cereale L.) were examined for five years in a field trial. In mid-June above-ground biomass of winter crops was removed and maize transplanted. The winter crops achieved maximum dry matter yield about three to five weeks before maturity. Mixed stands yielded more biomass than pure stands and exhibited greater yield stability. The relative advantage of intercropping, expressed as land equivalent ratio (LER), determined for intercropped winter pea/rye were 1.1 to 1.2 and for crimson clover/rye 1.3. At maturity, the amount of fixed nitrogen ranged between 178 kg N for crimson clover and 242 kg N ha^-1 for winter pea, respectively. At the end of anthesis (middle of June, harvesting stage for silage fodder) 75% and 88% of the total fixed nitrogen was achieved, for clover and pea, respectively. In intercropping the amount of fixed nitrogen was lower than in pure stands due to a lower seed density of the legume; however, the N-fixing efficiency was greater than in pure stands. N-release of the winter pea in a pure stand produced a maximum yield in maize (Zea mays L.) without additional N-fertiliser. An additional N mineral fertilisation of 75 to 150 kg N and 75 to 225 kg N was necessary to achieve maximum yields in maize following intercropped winter pea and crimson clover, respectively. Legumes in mixed stands with rye resulted in lower amounts of residual nitrogen after maize harvest. The beneficial effect of legumes on maize can be divided into N-effects and rotation effects. Both effects were positive regarding winter pea. The rotation effect of crimson clover in pure stands on maize was negative. Allelopathic effects and the high sensitivity of crimson clover to mineral nitrogen in the soil, released by residues of the preceding crop, winter rape (Brassica rapa L.), were discussed as the reason for this observation. The combination of the winter pea in pure stand and maize achieved the highest total biomass yield from winter and summer crops, unfertilised (156 dt ha^-1 dry). The combinations of intercropped legumes and maize produced biomass yields of 142 to 145 dt ha^-1. Because winter pea is highly susceptible to lodging, intercropping with low seed density of rye is recommended (3/4 winter pea, 1/4 rye). The rye crop prevents lodging by providing support and high rates of N-fixation are achieved with high seed density of pea. Intercropping with crimson clover and rye should be based on high seed densities of legumes, too because rye is highly competitive within those mixtures. This revised version was published online in August 2006 with corrections to the Cover Date.