Development of mass trapping technique for control of brinjal shoot and fruit borer, Leucinodes orbonalis (Lepidoptera: Pyralidae)

Locally-produced clear plastic water traps (12 cm x 14 cm base and 21 cm height) were optimized for use in large-scale mass trapping trials for control of brinjal fruit and shoot borer, Leucinodes orbonalis Guenée. Changing the shape (square and triangular) and number (two and four) of entry holes in the water trap had no significant effect on trap catch. Significantly more male moths were caught in traps treated with water containing powdered detergent than liquid detergent, light gear oil or insecticide. All water traps tested caught significantly higher numbers of moths than sticky delta traps with open sides under farmers' field conditions. Trap catches per 100 m2 were found to increase with increasing number of traps from 3 to 6 but the difference in catch between 4 and 6 traps per 100 m2 was not significant. Two small-scale replicated integrated pest management (IPM) trials were conducted consisting of the optimized water trap placed out with 10 m spacing (4 per 100 m2) and infested shoots pruned and destroyed. The first season trial had two treatments, IPM and farmers' practice in which farmers applied insecticide every two days in the peak harvest period. Overall, the percentage of healthy fruit and yields in both treatments were comparable at 53.8 and 49.6% and 20 and 19.4 tonnes per ha in the IPM and farmers' practice plots respectively. However, the initial infestations in the IPM plots (68%) were significantly higher than in farmers' practice plots (16%) due to the proximity of the nurseries used for the IPM plots to stacks of brinjal crop residues from the previous season that acted as a source of infestation. The second season's trials contained a third treatment in which IPM and farmers' practice were combined. The percent total healthy fruits harvested were 46.1, 58.6 and 69.1% respectively for the farmers' practice, farmers' practice plus IPM and IPM alone. Averaged total fruit yields were approximately 12 tonnes per ha for the farmers' practice plots and 30 tonnes per ha for each of the IPM-treated plots. The IPM plot had significantly fewer infested fruit than the IPM plus farmers practice plots and this was attributed to the activity of the larval parasitoid Trathala flavo-orbitalis (Cameron) that was suppressed in trial plots treated with insecticides.     

Female sex pheromone of brinjal fruit and shoot borer,Leucinodes orbonalis (Lepidoptera: Pyralidae): trap optimization and application in IPM trials

Delta and wing traps baited with synthetic female sex pheromone of Leucinodes orbonalis Guenée were found to catch and retain ten times more moths than either Spodoptera or uni-trap designs. Locally-produced water and funnel traps were as effective as delta traps, although 'windows' cut in the side panels of delta traps significantly increased trap catch from 0.4 to 2.3 moths per trap per night. Trap catch was found to be proportional to the radius of sticky disc traps in the range 5-20 cm radius, discs with a 2.5 cm radius caught no moths. Wing traps placed at crop height caught significantly more moths than traps placed 0.5 m above or below the crop canopy. Replicated integrated pest management (IPM) trials (3 x 0.5 ha per treatment) were conducted in farmers fields with young and mature eggplant crops. Farmers applied insecticides at least three times a week in all check and IPM plots. In addition pheromone traps were placed out at a density of 100 per ha and infested shoots removed weekly in the 0.5 ha IPM plots. Pheromone trap catches were reduced significantly from 2.0 to 0.4 moths per trap per night respectively in check and IPM plots in a young crop and 1.1 to 0.3 moths per trap per night in check and IPM plots respectively in a mature crop. Fruit damage was significantly reduced from an average of 41.8% and 51.2% in check plots of young and mature crops respectively to 22% and 26.4 respectively in the associated IPM plots. Significant differences in pheromone trap catches and fruit damage were attained four and two weeks respectively after IPM treatments began in the mature crop whereas in the immature crop significant differences were not observed for the first eight to nine weeks respectively. The relative impact of removing infested shoots and mass trapping on L. orbonalis larval populations was not established in these trials but in both cases there was an estimated increase of approximately 50% in marketable fruit obtained by the combination of control techniques compared to insecticide treatment alone.     

Volatiles from host plant brinjal attract the brinjal Fruit and Shoot Borer -Leucinodes orbonalis Guenee

Brinjal Fruit and Shoot Borer- Leucinodes orbonalis Guenee is a major insect pest on brinjal- Solanum melongena worldwide. An effective strategy used in developing pest controlling agents is the synergism between insect pheromones and host plant volatiles, which can increase the attraction of insect pest. The present study was aimed at investigating the chemical constituents and attractant effects of the volatiles extracted from different parts of the host plant brinjal on the behavior of adult L. orbonalis. Bioassay using Y-shaped olfactometer revealed that the one-day old virgin female, gravid female and male insects respond positively to the host plant volatiles extracted from fruits, leaves and shoots but not to that of flowers. It was shown that the gravid females were significantly attracted to all three volatiles (p < 0.05). Bioassay using X-shaped olfactometer identified that all three types of insects highly preferred the volatiles from fruits (p < 0.05). Gas chromatography-mass spectrometry analysis of volatiles indicated that brinjal plant produces volatile secondary metabolites, which include 2,2′-(Ethane-1,2-diylbis(oxy))bis(ethane-2,1-diyl) dibenzoate (12.11%), 3,7-dimethylocta-1,6-dien-3-ol (22.38%), Benzyl alcohol (22.9%) and Benzyl alcohol (27.06%) as major constituents from fruits, shoots, leaves and flowers respectively. Responses of insects to the volatiles from host plant in the absence of visual cues direct us to focus on the importance of host plant volatiles to locate the plant. Results of this study emphasize the major role that host plant volatiles play in the attraction of insect pests towards the plant.     

Proteome analysis of male accessory gland secretions in Leucinodes orbonalis Guenee (Lepidoptera: Crambidae), a Solanum melongena L. pest

Male accessory gland (MAG) proteins are transferred along with the sperm to females at the time of mating and have diverse effects on female reproductive physiology in a wide range of insects. In this study, we sought to identify the MAG proteins in Leucinodes orbonalis Guenee, a Solanum melongena L. pest, by analyzing the MAG proteins of virgin and mated male moths by nano-LC-ESI-MS/MS techniques. A total of 142 and 131 proteins in virgin and mated males were identified, respectively, among which 17 (12.0%) and 10 (7.6%) proteins were found to show secretory signals in virgin and mated males, respectively. These secretory proteins were shown to be involved in several biological processes in insects, including egg development, sperm-related functions/capacitation, defense, metabolism, and protein chaperoning. To the best of our knowledge, this is the first study to perform a proteome analysis of the MAG proteins of L. orbonalis, and offers an opportunity for further investigation of the functions of these proteins. In insects, certain MAG proteins are known to inhibit mating whereas others accelerate egg-laying. Therefore, the identification of these proteins in L. orbonalis may be useful for pest control.     

The role of intercrops of eggplant and cowpea on the control of Leucinodes orbonalis guenee (Lepidoptera: Crambidae)

BioControl - The eggplant fruit and shoot borer (EFSB), Leucinodes orbonalis Guenee (Lepidoptera: Crambidae) is the major pest of eggplant, Solanum melongena L., in Iran. Influences of...     

利用微卫星标记初步分析橘小实蝇4个地理种群的遗传多态性

橘小实蝇Bactrocera dorsalis(Bactrocera)(Hendel)是我国重要的检疫性害虫,对果蔬生产及其国际贸易造成很大影响。作者运用微卫星分子标记技术,用6对微卫星引物对采自我国福建、广东、云南3个地区及邻近国越南橘小实蝇种群间的遗传关系进行了初步分析,结果显示上述4个橘小实蝇地理种群间存在一定的遗传差异,地理隔离是造成这一差异的主要原因。     

AFLP genetic polymorphism in wild barley (Hordeum spontaneum) populations in Israel

The genetic diversity produced by the amplified fragment length polymorphism (AFLP) method was studied in 94 genotypes of wild barley, Hordeum spontaneum (C. Koch) Thell., originating from ten ecologically and geographically different locations in Israel. Eight primer pairs produced 204 discernible loci of which 189 (93%) were polymorphic. Each genotype had a unique banding profile and the genetic similarity coefficient varied between 0.74 and 0.98. The phenogram generated from these similarities by the UPGMA method did not group genotypes strictly according to their geographical origin, which pattern was also seen in the principal coordinate (PCO) plot. Genetic diversity was larger within (69%) than among (31%) populations. Associations between ecogeographical variables and the mean gene diversity were found at one primer pair. The results are discussed and compared with data obtained by the simple sequence repeat (SSR) method.     

Microsatellite polymorphism and genetic differentiation in three Norwegian populations of Elymus alaskanus (Poaceae)

 Variation at seven microsatellite loci was investigated in three local E. alaskanus populations from Norway and microsatellite variation was compared with allozyme variation. The percentage of polymorphic loci was 81%, the mean number of alleles per polymorphic locus was 5.7 and expected heterozygosity was 0.37. An F-statistic analysis revealed an overall 48% deficit of heterozygotes over Hardy-Weinberg expectations. Gene diversity is mainly explained by the within population component. The averaged between population differentiation coefficient, F _st , over 7 loci is only 0.13, which accounts for only 13% of the whole diversity and was contrary to allozyme analysis. The mean genetic distance between populations was 0.12. However, a χ² -test showed that allele frequencies were different (p < 0.05) among the populations at 5 of the 7 loci. In comparison with the genetic variation detected by allozymes, microsatellite loci showed higher levels of genetic variation. Microsatellite analysis revealed that population H10576 possesses the lowest genetic variation among the tested three populations, which concur with allozyme analysis. The dendrogram generated by microsatellites agreed very well with allozymic data. Our results suggest that natural selection may be an important factor in shaping the genetic diversity in these three local E. alaskanus populations. Possible explanations for deficit heterozygosity and incongruence between microsatellites and allozymes are discussed. Received November 6, 2001; accepted April 24, 2002 Published online: November 14, 2002 Addresses of the authors: Genlou Sun (e-mail: Genlou.sun@STMARYS.CA), Biology Department, Saint Mary's University, Halifax. Nova Scotia, B3H 3C3, Canada. B. Salomon, R. von Bothmer, Department of Crop Science, The Swedish University of Agricultural Sciences, P.O. Box 44, SE-230 53, Alnarp, Sweden.     

Host-plant-associated genetic differentiation in Northern French populations of the European corn borer

The phytophagous insects that damage crops are often polyphagous, feeding on several types of crop and on weeds. The refuges constituted by noncrop host plants may be useful in managing the evolution in pest species of resistance to the Bacillus thuringiensis toxins produced by transgenic crops. However, the benefits of these refuges may be limited because host-plant diversity may drive genetic divergence and possibly even host-plant-mediated sympatric speciation. The European corn borer, Ostrinia nubilalis Hübner (Lepidoptera: Crambidae), is the main pest of maize in Europe and North America, where it was introduced early in the 20th century. It has a wide host range but feeds principally on mugwort (Artemisia vulgaris L.) and maize (Zea mays L.). O. nubilalis is found on mugwort only in the northern part of France, whereas it is found on maize throughout France. The extent of genetic variation at allozyme markers was investigated in populations collected from the two host plants over the entire geographical distribution of the European corn borer on mugwort in France. Allelic differentiation between pairs of populations and hierarchical analyses of pools of samples from each host plant indicate that the group of populations feeding on maize differed from the group of populations feeding on mugwort. Our results suggest (1) host-plant-related divergent selection at the genomic region surrounding the Mpi locus and (2) limited gene flow between the populations feeding on mugwort and those infesting maize fields. These data indicate that adults emerging from mugwort would not be useful for managing the evolution of resistance to the B. thuringiensis toxins in European corn borer populations.     

First report of a hymenopteran parasitoid complex on jackfruit shoot and fruit borer Diaphania caesalis (Lepidoptera: Crambidae) from India

ABSTRACT

A survey of the species composition and relative abundance of Diaphania caesalis Walker (Lepidoptera: Crambidae) in plantations of jackfruit (Artocarpus heterophyllus Lam.) in three southern states along India’s western coast recorded thirteen species of primary parasitoids and one species of a hyperparasitoid, namely Tetrastichus pantnagerensis Khan (Hymenoptera: Eulophidae). Apanteles stantoni Ashmead (Hymenoptera: Braconidae) was the most dominant parasitoid and found in 35% of the parasitised host larvae in Karnataka, 38.5% in Kerala, and 33.1% in Tamil Nadu whereas the overall level of parasitisation, involving all the different parasitoids recorded, was approximately 41%–45% and did not vary significantly with location.     

Effects of artificial shade on attack by the mahogany shoot borer, Hypsipyla robusta (Moore)

Abstract

• 1 Swietenia macrophylla King (Meliaceae: Swietenioideae) provides one of the premier timbers of the world. The mahogany shoot borer Hypsipyla robusta Moore (Lepidoptera: Pyralidae) is an economically important pest of S. macrophylla throughout Asia, Africa and the Pacific. No viable method of controlling this pest is known. Previous observations have suggested that the presence of overhead shade may reduce attack by H. robusta, but this has not been investigated experimentally. This research was therefore designed to assess the influence of light availability on shoot‐borer attack on S. macrophylla, by establishing seedlings under three different artificial shade regimes, then using these seedlings to test oviposition preference of adult moths, neonate larval survival and growth and development of shoot borer larvae.
• 2 Oviposition preference of shoot borer moths was tested on leaves from seedlings grown under artificial shade for 63 weeks. A significant difference in choice was recorded between treatments, with 27.4 ± 1.5 eggs laid under high shade and 87.1 ± 1.8 under low shade.
• 3 Neonate larval survival on early flushing leaflets of S. macrophylla did not differ significantly between shade treatments. Larval growth rate, estimated by measuring daily frass width, was significantly higher for those larvae fed on seedlings from the high and medium shade treatments (0.1 mm/day), than the low shade treatment (0.06 mm/day). In laboratory‐reared larvae, the total mass of frass produced was significantly higher in the high shade treatment (0.4 g) than under the low shade treatment (0.2 g).
• 4 Longer tunnel lengths were bored by larvae in plants grown under high shade (12.0 ± 2.4 cm) than under low shade (7.07 ± 1.9 cm). However, pupal mass under low shade was 48% higher than that under the high shade treatment, suggesting that plants grown under high shade were of lower nutritional quality for shoot borer larvae.
• 5 These results indicate that shading of mahogany seedlings may reduce the incidence of shoot borer attack, by influencing both oviposition and larval development. The establishment of mahogany under suitable shade regimes may therefore provide a basis for controlling shoot borer attack using silvicultural approaches.

     

Genetic variation in host susceptibility to attack by the mahogany shoot borer, Hypsipyla grandella (Zeller)

1 Attack by shoot borers (Hypsipyla grandella Zeller) is the main factor limiting the cultivation of Cedrela odorata and Swietenia macrophylla, two economically important members of the mahogany family. No viable methods of pest control are currently available. To assess for genetic variation in susceptibility to pest attack, a combined progeny/provenance test of C. odorata and a provenance test of S. macrophylla were established separately at CATIE, Turrialba, Costa Rica, and assessed intensively over an 84-week period. 2 Variation in height growth between provenances was highly significant in both species, provenance means varying by a factor of 2.7 and 1.2 in C. odorata and S. macrophylla, respectively. Cedrela odorata also displayed provenance variation in foliar phenology, as 94.7% of the trees from a ‘wet zone’ provenance remained foliated, whereas > 35% of trees from three ‘dry zone’ provenances abscised their leaves during the dry season. 3 Both species displayed significant genetic variation in susceptibility to shoot borer attack. At the peak of attack during the second year of growth, the effect of provenance was highly significant in both species, the mean number of attacks per tree varying between 0.8–2.4 and 0.6–1.3 in different provenances of C. odorata and S. macrophylla, respectively. A provenance of C. odorata from San Carlos, Costa Rica, displayed consistently lower susceptibility to pest attack, being subjected to fewer attacks during the first year and demonstrating a higher mean height to first point of damage. 4 The nitrogen, total tannin and proanthocyanidin concentration of foliage varied significantly between C. odorata provenances: nitrogen concentration was significantly lower and tannin and proanthocyanidin contents were significantly higher in trees from the San Carlos provenance, Costa Rica. This study therefore suggests that proanthocyanidins may reduce susceptibility of C. odorata to H. grandella, at least during the early period of growth. 5 These results provide support for the development of plant resistance as a strategy for managing shoot borers. In particular, there may be scope for selecting for high foliar proanthocyanidin content and the ability to tolerate attack by vigorous apical growth.     

Relative Efficacy of Some Insecticides Against Brinjal Fruit and Shoot Borer,Leucinodes orbonalis Guen., and Their Impact on Fruit Yield

Six insecticides and their eight combinations were tested for their efficacy against brinjal fruit and shoot borer, Leucinodes orbonalis. Endosulfan + deltamethrin (0.07%, 0.0025%) and endosulfan + fenvalerate (0.07% + 0.005%) were highly effective against fruit borer that recorded only 13.3% damage as compared to 69.8% in control. The other promising treatments which significantly reduced the fruit damage over the control were in the order: carbaryl + fenvalerate = dichlorvos + fenvalerate (14.9%) > malathion + fenvalerate (16.4%) > fenvalerate + deltamethrin (16.6%) > dichlorvos = carbaryl + deltamethrin = malathion = dichlorvos + deltamethrin = malathion + deltamethrin (18.3%) > endosulfan (20.0%) > carbaryl (21.6%) with mean percentage of damage 14.9, 16.4, 18.3, 20.0, 21.6 and 69.8%, respectively. Carbaryl was least effective, but its combinations with pyrethroids were proved superior over carbaryl alone. Cost - benefit ratio ranged from a minimum of 1: 5.10 (carbaryl) to a maximum of 1: 20.44 (fenvalerate). Dichlorvos + fenvalerate combination gave the highest yield of 263.45 q/ha, whereas carbaryl was least effective giving 225.7 q/ha. with a net gain of Rupees 42,443.00 (US$ 886.00) and 28,141.00 (US$ 587.49), respectively. The other treatments were intermediate between the two insecticide regimes. However, all the treatments were superior over the control which produced 113.58 q/ha with a net gain of Rupees 340.00 only.     

Molecular genetic polymorphism of androgen receptor gene (AR) in African populations of Hadza and Datoga

The molecular genetic analysis of the polymorphic variants of the CAG repeat-containing locus of the androgen receptor (AR) gene was performed in the populations of Hadza and Datoga. Allele frequency distribution patterns were established. Alleles containing 20–25 repeats were the most abundant in both populations. The populations studied were compared with Asians (Han), white Americans, and Africans (Ariaal). Statistically significant difference between populations of Hadza and Datoga in the distribution of the AR allelic variants was demonstrated.     

Genetic control of fruit polymorphism in the GenusFedia (Valerianaceae) in the light of dimorphic and trimorphic populations ofF. pallescens

The genusFedia consists of three species (F. cornucopiae, F. graciliflora andF. pallescens) of winter annual herbs, endemic to the western Mediterranean Basin. The deciduous terminal fruits of these taxa are polymorphic in the development of their pericarp and/or calyx, and each population is dimorphic or more rarely trimorphic. The three main fruit types are dispersed in several manners, and are specialized for either epizoochory or myrmecochory. On the basis of our experimental study of dimorphic and trimorphic populations ofF. pallescens subsp.pallescens, a genetic model is presented in order to explain the control of this intrapopulational polymorphism. It is postulated that two diallelic loci are tightly linked on the same chromosome in a functional supergene. One allele of each locus displays a dominance effect in the heterozygous state. Within the four possible homologous allelic segments, only two are present in the dimorphic populations, three in the trimorphic ones, and are otherwise associated in diverse combinations in the remaining taxa of the genus. Similar examples of fruit polymorphism are already documented in the tribeValerianeae, subtribeFediinae. The hypothesis is put forward that this fruit polymorphism is a synapomorphy for the subtribe.     

Mitochondrial DNA polymorphism in three Antillean island populations of the fruit bat, Artibeus jamaicensis

The Neotropical fruit bat, Artibeus jamaicensis, occurs throughout Latin America and on many islands in the Caribbean. Populations from Jamaica (in the Greater Antilles) to Barbados (in the Lesser Antilles) have been classified as a subspecies (A.j. jamaicensis) separate from that on the Lesser Antillean island of St. Vincent (A.j. schwartzi). Mitochondrial DNA (mtDNA) was isolated from 54 individuals collected on these islands, analyzed by digestion with restriction endonucleases, and the restriction sites were mapped. Three different mtDNA genotypes (16,000 +/- 200 bp) were identified: J-1 (16 animals from Jamaica, one from St. Vincent, 15 from Barbados), J-2 (two animals from Jamaica), and SV-1 (18 animals from St. Vincent, two from Barbados). The J-1 and J-2 genotypes were estimated to differ from each other by only 0.4%, but the SV-1 genotype differed from J-1 and J-2 by 8.1%-10.5%. The estimated sequence divergence between SV-1 and J-1 is unusually large for mammals that are regarded as conspecific. Restriction mapping showed that the differences among the genotypes (presence or absence of particular restriction sites) were located throughout the genome. The presence of the J-1 mtDNA genotype on Jamaica and on St. Vincent and Barbados (1,400 km away) demonstrates that maternal lineages in these bats are not necessarily confined to single islands or limited geographic regions. The presence of the J-1 mtDNA genotype within the A. j. schwartzi population on St. Vincent and the presence of the SV-1 genotype in two specimens of A. j. jamaicensis from Barbados document genetic exchange between subspecific populations on these islands, which are separated by 180 km of open water.     

Allozyme differentiation among nine populations of the corn borer (Ostrinia) in China

To test the hypothesis of the migration of the corn borer, the allozymes of nine populations of the corn borer(Ostrinia) in China were checked using verticalslab polyacrylamide gel electrophoresis. Eight loci of six allozymes were analyzed. The mean of the genetic identities among the nine populations calculated from the allele frequencies was 0.99068, much closer than that of other species and geographical populations. The mean (0.97955) of the genetic identities between the XJYN population (Ostrinia nubilalis Hübner, collected from Yining, Xinjiang Autonomous Region) and each of the other eight populations (O. furnacalis Guenée) was significantly smaller than that between the pairs of the eight populations (0.99386;t test,P<0.01). Although the population XJYN clearly deviates from the other eight populations in the dendrogram, the relationship of the two species of corn borer was very close. It is possible that the speciation of corn borer may have resulted from single-gene substitutions.     

Insecticidal activity of Citrus aurantium fruit, leaf, and shoot extracts against adult olive fruit flies (Diptera: Tephritidae)

Solvent extracts of differing polarity from Citrus aurantium (L.) (Rutaceae) fruit, leaves, and shoots were evaluated for biological activity against adults of the olive fruit fly, Bactrocera oleae (Gmelin) (Diptera: Tephritidae). Using a petri dish residual exposure bioassay, we found that the petroleum ether extract from fruit alone showed insecticidal activity against the flies. The extract of the three fruit tissues (flavedo [peel], albedo, and flesh) indicated that bioactivity was limited to the flavedo, and this activity was significantly higher than that of the whole fruit extract. The most effective extract was obtained when fresh flavedo was used, whereas extracts of oven-dried flavedo were inactive. Fruit maturity also affected bioactivity; extracts of ripe fruit were more effective than those of unripe fruit. Our results suggest that C. aurantium flavedo contains secondary metabolites with insecticidal activity against B. oleae adults.     

High degree of genetic polymorphism in apolipoprotein(a) associated with plasma lipoprotein(a) levels in Japanese and Chinese populations

We have developed a sensitve, high-resolution method for the analysis of the apolipoprotein(a) [apo(a)] isoforms using sodium dodecyl sulfate (SDS)-agarose/ gradient polyacrylamide gel electrophoresis. In an analysis of the genetic polymorphism of apo(a) isoforms and their relationship with plasma lipoprotein(a) [Lp(a)] levels in Japanese and Chinese, this method identified 25 different apo(a) isoforms and detected one or two apo(a) isoforms in more than 99.5% of the individuals tested. The apparent molecular weights of the apo(a) isoforms ranged from 370 kDa to 950 kDa, and 22 of the 25 different apo(a) isoforns had a higher molecular weight than of apo B-100. Studies on Japanese families confirmed the autosomal codominant segregation of apo(a) isoforms and the existence of a null allele at the apo(a) locus. The observed frequency distribution of apo(a) isoform phenotypes fit the expectations of the Hardy-Weinberg equilibrium in both the Japanese and Chinese populations. Our data indicate the existence of at least 26 alleles, including a null allele, at the apo(a) locus. The frequency distribution patterns of the apo(a) isoform alleles in Japanese and Chinese were similar to each other and also similar to that of apo(a) gene sizes reported in Caucasian American individuals. The average heterozygosity at the apo(a) locus was 92% in Japanese and 93% in Chinese. A highly significant inverse correlation was observed between plasma Lp(a) levels and the size of apo(a) isoforms in both the Japanese (r=-0.677, P=0.0001) and the Chinese (r=-0.703, P=0.0001). A highly skewed distribution of Lp(a) concentrations towards lower levels in the Japanese population may be explained by high frequencies of alleles encoding large apo(a) isoforms and the null allele.