Mating behavior and male mating success in wildAnastrepha Ludens (Diptera: Tephritidae) on a field-caged host tree

Mating behavior and factors affecting mating success of males were studied using wild Anastrepha ludens on a fieldcaged host tree. The most common courtship sequence had five components: (1) male calls from the underside of a leaf, (2) female arrives to the maleoccupied leaf, (3) male orients to female and stops calling, (4) one or both approach to a face-to-face position 1–3 cm apart, and (5) male mounts female after 1–2 s. Courtship behavior was almost identical to that of laboratoryculture flies observed previously under laboratory conditions. Most malefemale encounters occurred at a height of 1–2m, well inside the outer canopy of the tree. Differential mating success by males occurred. No male mated more than once per day, owing possibly to a very short sexual activity period. Factors favoring mating success of males were survival ability and tendency to join male aggregations and to fight other males. Thorax length and age (9–11 days difference) had no effects on male copulatory success. Overall win/loss percentage was not related to mating success because the males that were most successful at mating fought mostly among themselves, driving their win/loss percentage down. However, these successful males (at mating) won most of their fights against less successful males. Results confirmed a lek mating system: males aggregated, called, and defended territories; territories did not contain femalerequired resources; and females exercised mate choice, apparently through selection of sites within leks.     

Starvation and the Mating Success of Wild Male Mediterranean Fruit Flies (Diptera: Tephritidae)

Recent evidence suggests that the nutritional state of male Mediterranean fruit flies, Ceratitis capitata (Wied.) (medfly), is an important influence on various components of their reproductive biology, including mating success. The objective of the present study was to examine experimentally the effect of temporary starvation on the mating success of wild male C. capitata. Males were maintained on protein–sugar or sugar-only diets, and for each diet we compared the mating success of continuously fed males versus males starved for 18 or 24 h immediately before testing. In trials conducted on field-caged, host trees, males starved for 24 h obtained only about half as many matings as fed males for both diets. However, when the starvation period was 18 h, starved males reared on the protein–sugar diet mated significantly less frequently than fed males, whereas starved males reared on sugar mated as often as fed males. Measurements of male pheromone calling and female attraction revealed that reduced mating success likely reflected the decreased signaling activity of starved males.     

Two-way selection for mating activity in the Mediterranean fruit fly, Ceratitis capitata

No significant variation in mating activity was observed among eight laboratory strains of the Mediterranean fruit fly (medfly), Ceratitis capitata (Wiedemann) (Diptera: Tephritidae). Two methods were used to select strains showing high and low mating activity: a single pair technique (SP) assessing time to mating and a mass technique (M) based on a mating index. Reciprocal pairings between fast and slow selected lines showed that the difference between the SP lines was female-determined whereas the difference between the M lines depended on the behaviour of both sexes. M selection, irrespective of its direction, affected two courtship parameters, vibration distance and vibration duration both of which tended to be shorter. M selection was also associated with a reduction in startle activity in females.

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Résumé Aucune variation significative n'a été observée dans les activités sexuelles de huit souches de laboratoire de la mouche méditerranéenne du fruit, Ceratitis capitata (Wiedemann) (Diptera Tephritidae).Deux méthodes ont été utilisées pour sélectionner des souches manifestant une activité sexuelle basse ou élevée: une technique utilisant un seul couple (SP), basée sur la durée de la période avant copulation, et une technique de masse (M), basée sur un index d'accouplement.Des appariements réciproques entre les lignées rapides et lentes ont montré que la différence entre les lignées SP était déterminée par les femelles, tandis que la différence entre les lignées M résultait du comportement des deux sexes. La sélection M, quelle que soit sa direction, affects deux paramètres, la distance entre partenaires lors de la vibration et la durée de la vibration qui tous deux tendent à se raccourcir. La sélection M est aussi associée à une réduction de la réaction des femelles à un brusque stimulus lumineux (Startle activity).

     

Mitochondrial DNA restriction map for the mediterranean fruit fly,Ceratitis capitata

Molecular genetic research on the Mediterranean fruit fly,Ceratitis capitata, will provide tools to permit determination of source populations for new pest infestations. Restriction fragment length polymorphism (RFLP) of mitochondrial DNA provides some interpopulation discrimination. A restriction map, including the informative variableEcoRV andXbaI restriction sites, is constructed for the Mediterranean fruit fly, and several restriction sites are associated with specific gene regions based on polymerase chain reaction-RFLP and sequence analyses. A partial sequence of the mitochondrial 16S ribosomal RNA gene is reported.     

Host-marking pheromone and use of previously established oviposition sites by the mediterranean fruit fly (Diptera: Tephritidae)

Under controlled conditions, the Mediterranean fruit fly, Ceratitis capitata(Wiedemann) preferred to initiate oviposition into preexisting, naturally formed oviposition punctures in a host fruit, kumquat (Fortunella japonica),over establishing new sites on the fruit. This preference was expressed despite the presence of naturally deposited host-marking pheromone (HMP)shown previously to deter oviposition. An almost-identical preference for existing punctures was expressed when females were presented with fruit bearing artificially made punctures on which HMP had been naturally deposited. Using artificial punctures and HMP extracts, the occurrence of punctures was manipulated independently of the presence of HMP.Under field-cage conditions, we found that (1) punctures stimulated egg-laying on kumquats, regardless of HMP treatment; (2) HMP extract inhibited egg-laying, regardless of the occurrence of punctures; and (3) the extent to which HMP inhibited egg-laying was greater on fruit free of punctures than on fruit bearing them. The physiological, evolutionary, and pest management implications of these results are discussed.     

Effect of pupation environment on the quality of pupae and adults of the Mediterranean fruit fly

When mature larvae of the Mediterranean fruit fly, Ceratitis capitata (Wiedemann) pupated in bran rather than naked there was an increase in adult eclosion, but no difference in 9 day pupal weight, 9 day lipid content, or flight ability. There was no difference when pupation occurred at 20 or 25°C. Pupation in sand at 20°C caused severe mortality, while pupation in sand at 25°C did not. Data are presented which favour naked pupation over pupation in any medium, and evidence is presented which indicates that collection of mature larvae in a rotary tumbler rather than by the conventional popping method is economically preferable and is without significant biologically adverse effects.

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Résumé Si des larves mûres de Ceratitis capitata se nymphosent dans du son ou à l'air libre, dans le premier cas le taux d'éclosions est plus élevé, mais le poids et la teneur en lipides au neuvième jour de nymphose ainsi que l'aptitude au vol sont identiques dans les deux. Il n'y a pas de différences pour les pupaisons à 20°C et 25°C, mais la pupaison dans le sable à 20°C provoque une forte mortalité, tandis qu'à 25°C le sable n'a aucun effet. Les résultats obtenus sont favorable à une pupaison à nu plutôt que dans un substrat, et des arguments sont apportés montrant que la collecte des larves mûres dans un culbuteur rotatif ne provoque significativement pas plus d'effets nocifs que la méthode classique du saut.

     

Effect of larval host on life history traits of the mediterranean fruit fly,Ceratitis capitata

Summary Demographic parameters for the Mediterranean fruit fly reared on each of twenty four different hosts from sixteen different plant families are reported. These include cohort parameters of development, survival, pupal sizc, and fecundity as well as population parameters such as intrinsic rate of population increase (r) and mean generation time. Major findings include the following: i) no consistent quantitative relationships existed between r and its chief demographic determinants such as preadult developmental time and adult fecundity; and ii) few correlations existed among the cohort life history parameters themselves. The principle conclusion is that the medfly is a successful generalist frugivore because of its developmental ability to offset the effect of the value for a host-specific trait that tends to lower r with one that tends to increase r, the net result of which is to maintain a relatively high r.     

Attraction of female Chiracanthium mildei(Araneae: Clubionidae) to olfactory cues from maleMediterranean fruit flies Ceratitis capitata(Diptera: Tephritidae)

Chiracanthium mildei, a common sac spider (Clubionidae) in Israel, is an important predator of pest insects in citrus and apple orchards. Field observations indicate that the females of this hunting spider are attracted to traps containing the parapheromone trimedlure and males of the Mediterranean fruit fly (Ceratitis capitata),which were caught in these traps. Laboratory testsshow that C. mildei females are attractedto olfactory cues from male C. capitata, but notto trimedlure. This suggests that C. mildeimay be an important nocturnal predator of C.capitata.     

Influence of male aggregation size on female visitation in Bactrocera tryoni (Froggatt) (Diptera: Tephritidae)

Abstract  The importance of male aggregation size for female visitation and initiation of male pheromone-calling was investigated in Bactrocera tryoni (Froggatt) (Diptera: Tephritidae) using artificial male aggregations in large laboratory cages. Female B. tryoni visited the largest aggregation more frequently than single males in association with a higher proportion of calling males, but there was no correlation between aggregation size and female visitation. Female B. tryoni had a limited capacity to perceive a difference between the number of calling males. Calling propensity of male B. tryoni was increased by the presence of conspecific males. Increased calling propensity in larger groups of male B. tryoni may be due to social facilitation of male calling behaviour. Female visitation at aggregations was only weakly associated with male calling, suggesting that aggregation size and the number of pheromone-calling males are not the only factors important in locating mates in B. tryoni , and it is possible that low-density populations could persist so long as females can encounter single males.     

Olfactory and Behavioral Mechanisms Underlying Enhanced Mating Competitiveness Following Exposure to Ginger Root Oil and Orange Oil in Males of the Mediterranean Fruit Fly, Ceratitis capitata (Diptera: Tephritidae)

Males of the Mediterranean fruit fly, Ceratitis capitata, are strongly attracted to various plant odors, and previous work has demonstrated that male exposure to certain odors, including the scent of orange oil (OO) and ginger root oil (GRO), increases their mating success relative to non-exposed males. However, the mechanism(s) underlying this mating increase is not known. Here, we describe several experiments that further investigate the association between GRO- and OO-exposure and male signaling activity, pheromone attractiveness, and mating success in male medflies. Exposure to GRO or OO increased time spent pheromone calling but did not accelerate the rate of male sexual maturation. Using a wind tunnel, we compared female attraction to the pheromone of control, non-exposed males versus males previously exposed to OO or GRO. There was no evidence that GRO exposure enhanced the attractiveness of the male pheromone. The data for OO were inconclusive: females tended to spend more time on spheres emanating pheromone from OO-exposed males than on spheres emanating pheromone from non-exposed males, but the number of female landings did not differ between the two types of pheromone sources. Female choice tests confirmed that GRO- and OO-exposure boost male mating success relative to non-exposed males. Application of GRO directly to the abdomen reduced male mating success, whereas similar application of OO boosted male mating success. The potential role and mode of action of plant chemicals in the mating behavior of male medflies are evaluated in light of these findings.     

Application of quality control procedures to large scale rearing of the Mediterranean fruit fly

Quality control charting was applied to a number of criteria implemented to monitor the large scale (ca 10 million pupae per week) production of Ceratitis capitata (Wiedemann) and to assess the quality of the flies so produced. The control chart procedure was useful and indicated the development involving adult flight ability. It also illustrated the variability inherent in some parameters e.g. percent recovery of pupae from eggs, but in the absence of similar data from other rearing facilities it is not possible to state whether or not that level of virability is inherent in the process.

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Résumé Une procédure de contrôle de la qualité a été appliquée à une série de critères de l'élevage de masse (c'est-à-dire 10 millions de pupes var semaine) de C. capitata pour garantir la qualité des mouches produites. La charte de contrôle a été utile et a permis de mettre en évidence un problème concernant l'apptitude au vol des adultes. Elle a illustré la variabilité de certains paramètres, par exemple le taux de pupes obtenues à partir des oeufs, mais en l'absence de données semblables provenant d'autres élevages, il n'est pas possible de savoir quelle part du taux de variabilité est inhérente ou non à la méthode.

     

Odour-mediated foraging by yellowjacket wasps (Hymenoptera: Vespidae): predation on leks of pheromone-calling Mediterranean fruit fly males (Diptera: Tephritidae)

Predation is probably the most important male mortality factor in insect species with courtship displays that render males performing them conspicuous targets of predators. Sexually active Mediterranean fruit fly males, Ceratitis capitata (Wied.), aggregate in leks, where they participate in agonistic encounters and engage in visual, acoustic and pheromone-calling displays to attract receptive females. The objective of this study was to assess: a) whether sexually displaying C. capitata males in leks inside host and non-host foliage are subject to predation by the most prominent predators yellow-jacket wasps, Vespula germanica (F.), and if so, b) whether olfactory, visual or auditive stimuli are used by foraging wasps in locating male C. capitata prey. Studies were carried out in a citrus orchard and surroundings on the island of Chios, Greece. Observations were conducted using perforated containers hung within mulberry, fig or citrus foliage. Living C. capitata flies of different sex and either mature or immature were placed inside. Our results show that the yellowjacket wasps have learned to associate the presence of sexually active medfly males aggregated in leks with their prey's pheromone (kairomone). Foraging wasps, flying through the crowns of host trees, responded to the odour source of C. capitata male pheromone by approaching from downwind. Even inside dense citrus tree foliage, wasps keyed in on aggregations of pheromone-calling males using olfactory stimuli. Stimuli of visual and acoustic male signalling were only used at close range, after having followed the pheromone plume close to its source. Visual cues played a greater role in directing wasp foraging under more open and exposed host foliage conditions. Odour-based foraging of wasps inside host foliage in the mid-morning hours, when medfly male lekking activities peak, shifted gradually to a more visual-based host fruit patrolling in the afternoons to capture ovipositing and feeding medfly females. On ripe fruit, particularly fig, V. germanica visual prey hunting also included the capture of feeding medfly males, other feeding Diptera, as well as medfly larvae extracted from wasp-made perforations in the fruit.     

Population genetics of the potentially invasive African fruit fly species, Ceratitis rosa and Ceratitis fasciventris (Diptera: Tephritidae)

A set of 10 microsatellite markers was used to survey the levels of genetic variability and to analyse the genetic aspects of the population dynamics of two potentially invasive pest fruit fly species, Ceratitis rosa and C. fasciventris, in Africa. The loci were derived from the closely related species, C. capitata. The degree of microsatellite polymorphism in C. rosa and C. fasciventris was extensive and comparable to that of C. capitata. In C. rosa, the evolution of microsatellite polymorphism in its distribution area reflects the colonization history of this species. The mainland populations are more polymorphic than the island populations. Low levels of differentiation were found within the Africa mainland area, while greater levels of differentiation affect the islands. Ceratitis fasciventris is a central-east African species. The microsatellite data over the Uganda/Kenya spatial scale suggest a recent expansion and possibly continuing gene flow within this area. The microsatellite variability data from C. rosa and C. fasciventris, together with those of C. capitata, support the hypothesis of an east African origin of the Ceratitis spp.     

The role of ammonia in the attraction of females of the Mediterranean fruit fly to protein hydrolysate baits

Attraction of female Mediterranean fruifly (medfly) to ammonia and to protein baits was studied with an olfactometer. Ammonia bait (1 cc/trap) proved to be an effective lure. The maximum number of females was caught in traps loaded with 0.01 M ammonia solution with a release rate of 5.28 g/cc/h.A positive correlation was found between female catch and the ammonia release rate of various protein baits, especially from dry protein hydrolysate of which casein hydrolysate was the most effective. However, casein hydrolysate was less attractive than certain ammonia solutions having a lower rate of ammonia release. Elevation of the pH of the liquid commercial baits, Buminal and Naziman, increased the latter's efficacy as medfly baits but the increased stimulation could not be strictly correlated with the increased rate of ammonia release. It is therefore suggested that medfly olfactory response to protein baits is affected not only by ammonia but by other volatiles as well.

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Résumé L'attraction de la femelle de la mouche méditeranéenne à l'ammoniaque et aux attractifs protéiniques a été étudiée au moyen de l'olfactomètre décrit par Gothilf & Galun (1982). L'ammoniaque (1 cc/piege) s'avéra être un attractif efficace. Le nombre maximum de femelles fut attrappé dans les pièges chargés avec une solution d'ammoniaque 0.01 M ayant une vitesse de diffusion de 5.28/g/cc/h.Une corrélation positive a été trouvée entre les captures de femelles et les vitesses de diffusion de divers attractifs protéiniques, particulierement ceux à base d'hydrolysats de protéines secs parmi lesquels le plus effectif a été l'hydrolysat de caséine. Toutefois, l'hydrolysat de caséine a été moins attractif que certaines solutions d'ammoniaque ayant une vitesse de diffusion plus basse. Une hausse du pH des attractifs liquides commerciaux, Buminal et Naziman, a augmenté leur efficacité en tant qu'attractifs pour la mouche méditeranéenne; toutefois la stimulation plus intense n'a pu étre mise en correlation stricte avec une vitesse accrue de la diffusion de l'ammoniac. C'est pourquoi il est suggéré que la réaction olfactive de la mouche méditerranéenne est régie, non seulement par l'ammoniaque mais aussi par d'autres substances volatiles.

     

Parasitism of fruit fliesCeratitis capitata andAnastrepha SPP. [Diptera: Tephritidae] in Guatemala

In a 4-year surveyDoryctobracon crawfordi Viereck andBiosteres longicaudatus Ashmead were the most common parasitoids recovered from bothCeratitis capitata (Wiedemann) andAnastrepha spp. Other braconids recovered from both genera wereD. areolatus Szepligeti,Bracanastrepha anastrephae (Viereck), and a Cynipid. Combined parasitization in both generally by ranged from 0.04 to 7.95% (Av. 1.5%). The numbers were particularly low in economically important fruit crops such as coffee, orange, and peach. Combined parasitoid population was correlated with the combined fruit fly infestation at 5 elevation strata.      

Assessment of quality control parameters during mass rearing of a pupal colour genetic sexing strain of the Mediterranean fruit fly

Quality control charts were employed to monitor the quality of several parameters during mass rearing of the T : Y(wp ⁺)30C strain of Ceratitis capitata (Wiedemann), in which male and female flies emerge from brown and white puparia, respectively. Egg hatch and pupal recovery in this genetic sexing strain were below that reported for a wild-type Sohag strain, whereas pupal size, adult emergence and flight ability were higher. The overall recovery from eggs to adult flies capable of flight was similar in the two strains but egg production in T:Y(wp ⁺)30C was reduced to about 56% of that of Sohag. The quality control charts showed an acceptable level of variation within sets of data but the level between data sets was occasionally out of control. This indicated the presence of uncontrolled conditions during the rearing process, possibly arising from variations in the quality of the larval diet. It is concluded that the quality of T : Y(wp ⁺)30C renders this strain suitable for mass rearing in sterile insect technique (SIT) programmes requiring the elimination of the female flies prior to the release of sterilized males. The next step should involve further quality assessments both under field cage and under field release conditions. The importance of optimal and standardized dietary conditions is stressed in relation to the quality of mass reared medflies.

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Résumé Des normes de qualité ont été utilisées pour relever la valuer de plusieurs paramètres lors de l'élevage industriel de la souche T : Y(wp ⁺)30C de C. capitata Wied. dont les mâles et les femelles émergent respectivement de pupes brunes et blanches. Les taux d'éclosion et d'obtention de pupes sont plus faibles avec cette souche qu'avec la souche souvage Sohag; par contre, la taille des pupes, le taux d'émergence des adultes et l'aptitude au vol étaient plus élevés. La fréquence totale des adultes capables de voler par rapport au nombre initial d'ufs a été de 0,297 contre environ 0,311 pour Sohag. Cependant, la production d'ufs a été réduite à environ 56% de celle de Sohag. Les tableaux de contrôle de la qualité ont révélé un niveau de variation acceptable à l'interieur rieur des lots de données, mais la variabilité entre les lots excédait les normes admissibles. Ceci implique des paramètres incontrôlés pendant l'élevage de masse, probablement liés à des variations de la qualité de l'aliment. On peut en conclure que la qualité de la souche T : Y(wp ⁺)30C la rend utilisable pour un élevage de masse dans des programmes d'utilisation d'insectes stériles (SIT) exigeant l'élimination des femelles avant le lâcher de mâles stériles. La prochaine étape impliquera de nouvelles épreuves de qualité dans des cages en champ et dans les conditions de libération dans la nature. L'importance des conditions optimales d'une alimentation standardisée est soulignée par la qualité des mouches obtenues en élevage de masse.

     

Life history response of female Mediterranean fruit flies, Ceratitis capitata, to periods of host deprivation

Female medflies were subjected to four different patterns of host deprivation at each of three levels for a 24-day period. Treatments ranged from host absence 8 days post-eclosion to host absence 2 out of 3 days. Survival was recorded daily and egg production was recorded on the days in which hosts were present.The direct effect of host deprivation is to deny females the opportunity to oviposit. This disrupts their reproductive cycle and reduces their overall reproductive effort. Reduced reproductive effort at young ages increases survival (up to 2-fold) and daily reproduction (up to 8-fold) at older ages. Therefore, the long term effect of host deprivation is to postpone senescence.The implications of these findings regarding medfly sensescence are discussed in relation to: (i) reduction in population extinction rates during periods of host scarcity and (ii) egging strategies in medfly mass-rearing.

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Résumé Les femelles de C. capitata ont été privées d'hôtes suivant 4 modalités pendant 3 durées (33, 50 et 67% de leur vie) par période de 24 heures. Les modalités se répartissaient de la privation pendant 8 jours après l'éclosion jusqu'à 1 jour sur 3. La survie a été notée chaque jour et la ponte pour tous les jours où il y avait des hôtes.L'effect direct de l'absence d'hôte est d'empêcher la femelle de pondre. Ceci interrompt leur cycle de ponte et réduit leur activité reproductrice totale. Un effort reproductif réduit des femelles jeunes augmente la survie (jusqu'à 2 fois) et la ponte quotidienne (jusqu'à 8 fois) chez les femelles plus âgées. Ainsi, l'effet à long terme de la privation d'hôtes est d'ajourner la sénescence.Les implications de ces observations sur la sénescence de C. capitata sont discutées quant à: (1) la réduction des taux d'extinction des populations pendant les périodes de rareté d'hôtes; et: (2) les stratégies d'obtention de pontes dan les élevages de masse de C. capitata.

     

Pheromone deposition on leaf territories by male Caribbean fruit flies,Anastrepha suspensa (loew) (Diptera: Tephritidae)

Male Caribbean fruit flies apply everted anal membranes associated with pheromone glands to the substrate while on leaf territories. There is a peak in abdominal dipping at the onset of the photoperiod, followed by a decline and then a more extensive period of activity in the second half of the photoperiod. During peak signaling periods, most males had a distinct set of lateral abdominal phermone glands protruded for most of the time. However, the frequency of anal pheromone gland dipping varied considerably over time. Episodes of wing fanning (which may disperse pheromones) and abdomen dipping coincide during a male's tenure on a host leaf. Host-plant leaves were contained for 48 h with signaling males or immature females or kept without insects. Several, though not all, of the constituents of the pheromone were found on leaves within 1 h of removing signaling males. None of these compounds occurred on either type of control leaf. Mature virgin female flies were more likely to be found in contact with host leaves that had been previously exposed for 48 h to the activities of sexually mature males than with unexposed control leaves. There was no difference in the reaction of mature virgin female flies to unexposed control leaves or leaves previously exposed to the activities of other sexually immature female or male flies for 48 h. Presumably, the deposition of pheromones by mature males accounts for the difference in reactions.     

Sexual selection for symmetrical male medflies (Diptera: Tephritidae) confirmed in the field

The role of fluctuating asymmetry as an indicator of fitness to females in mate choice remains controversial. Previous studies indicated that male medflies with symmetrical supra-fronto-orbital (SFO) bristles achieve relatively high mating success under laboratory conditions. Here we present data from field cage studies of wild collected medflies in Guatemala and Crete, which reveal the same association between fluctuating asymmetry in SFO bristle length and mating success as that seen in the laboratory. The experiments in Crete included males that were missing one or both of their bristles. A comparison of mating success between the three groups indicated that the mere presence of bristles did not exert a major influence. Analysis of attempted courtships suggests that the association between male mating success and FA in bristle length appears to be generated as a result of females being more likely to enter into courtships with symmetrical males, rather than through a rejection of asymmetrical males during or after it. This raises the possibility that the primary stimulus that makes a symmetrical male attractive is acting at too great a distance to depend on symmetry itself. Alternatives might include superior pheromone emissions or the occupation of a prime location within the lek.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 81 , 347–355.