Development of polymorphic microsatellite markers in Hessian fly, Mayetiola destructor (Say)

A microsatellite library was prepared from size-selected genomic DNA of Hessian fly (Mayetiola destructor). Approximately 81% of recovered clones hybridized with microsatellite motif-specific probes. Subsequently, 2350 clones were sequenced. Sixty-two individual flies from laboratory strains were used to test for reliability and polymorphism in 50 of the microsatellites by gel electrophoresis; 18 were further tested with capillary electrophoresis. Of these, 17 behaved as a polymorphic single locus appropriate for population analysis.     

Distribution,population dynamics and damage of Hessian fly,Mayetiola destructor (Diptera: Cecidomyiidae) in North Tunisia

A three years survey and monitoring studies (2013–2014–2015) were carried out through 4 regions of north Tunisia in order to follow the evolution of the distribution, the frequency of occurrence and damage caused by the Hessian fly Mayetiola destructor (Say) to bread wheat (Triticum aestivum L.) and durum wheat (Triticum durum Desf). Moreover, the effectiveness of resistance genes H3, H5, H6, H7, H8, H9, H10, H11, H12, H13, H14, H15, H22, H23, H25 and H26 to protect wheat from Hessian fly attack was assessed in natural field and under controlled laboratory conditions at INRAT-Kef Station. Results showed that Hessian fly was detected in 60.33% and 51.5% of all sampled durum and bread wheat fields, respectively. This pest was more frequent with a higher percentage of infestation in semi-arid regions. Indeed, during 2013, infestation rate attained 12.39% in Kef region against 0.9% registered in Bizerte region. In order to update information about the annual number of generations, we surveyed the population dynamic of Hessian fly in Kef region. Three generations of the fly were counted annually on wheat, with two complete and one incomplete generation. This insect affects host plant growth at different developmental stages. Plant height was the most affected parameter followed by shoot dry weight and tiller number. Field investigations on host resistance revealed that among the 16 tested resistance genes, and only three were strictly effective (H22, H25 and H26). The resistance genes H5, H9, H13 and H9H13 have also conferred high levels of protection against Hessian fly. This work indicated that H22, H25 and H26 genes could be incorporated into Tunisian wheat varieties and released to farmers to manage the threat due to Hessian fly attacks.     

Feeding by Hessian fly [Mayetiola destructor (Say)] larvae does not induce plant indirect defences

Abstract.  1. Recent research has addressed the function of herbivore-induced plant volatiles in attracting natural enemies of feeding herbivores. While many types of insect herbivory appear to elicit volatile responses, those triggered by gall insects have received little attention. Previous work indicates that at least one gall insect species induces changes in host-plant volatiles, but no other studies appear to have addressed whether gall insects trigger plant indirect defences.
2. The volatile responses of wheat to feeding by larvae of the Hessian fly Mayetiola destructor (Say) (Diptera: Cecidomyiidae) were studied to further explore indirect responses of plants to feeding by gall insects. This specialist gall midge species did not elicit a detectable volatile response from wheat plants, whereas a generalist caterpillar triggered volatile release. Moreover, Hessian fly feeding altered volatile responses to subsequent caterpillar herbivory.
3. These results suggest that Hessian fly larvae exert a degree of control over the defensive responses of their host plants and offer insight into plant-gall insect interactions. Also, the failure of Hessian fly larvae to elicit an indirect defensive response from their host plants may help explain why natural enemies, which often rely on induced volatile cues, fail to inflict significant mortality on M. destructor populations in the field.     

Genes encoding a group of related small secreted proteins from the gut of Hessian fly larvae [Mayetiola destructor （Say）]

A group of related genes has been isolated and characterized from the gut of Hessian fly larvae [Mayetiola destructor （Say）]. Members in this group appear to encode proteins with secretary signal peptides at the N-terminals. The mature putative proteins are small, acidic proteins with calculated molecular masses of 14.5 to 15.3 kDa, and isoelectric points from 4.56 to 4.88. Northern blot analysis revealed that these genes are expressed predominantly in the gut of Hessian fly larvae and pupae. Two related genes, GIOK1 and GIOK2, were isolated as tandem repeats. Both genes contain three exons and two introns. The intron/exon boundaries were conserved in terms of amino acid encoding, suggesting that they arose by gene duplication. The fact that the frequency of this group of clones in a gut cDNA library higher than that of total cDNA clones encoding digestive enzymes suggested that this group of proteins may perform an important function in the gut physiology of this insect. However, the exact functions of these proteins are as yet known since no sequence similarity could be identified between these proteins and any known sequences in public databases using standard methods.     

Functional morphology of the dorsal vessel in the adult fly Protophormia terraenovae (diptera,calliphoridae)

The morphology and ultrastructure of the contractile tubular vessel acting as the cardiac pump in Protophormia terraenovae flies were analyzed by means of light microscopy, SEM and TEM. The results provide a novel anatomical picture of the two vessel portions, the abdominal heart and the aorta, and lay the foundations for an interpretation of the cardiocirculatory function in the fly. In the thorax, the thin and unchambered aorta is without apertures, while the abdominal heart presents a very small caudal aperture and pairs of lateral ostia, one in each of the five chambers of which it is composed. The ostia of the four more distal chambers are of the incurrent type, which is to say that they act as valves ensuring that hemolymph flows only into the heart. Conversely, the ostia in the most proximal chamber allow hemolymph to flow both into and out of the heart. The entire vessel is composed of a single layer of myofibers that are oriented circularly around the lumen in the abdominal heart and longitudinally in the thoracic aorta. The abdominal heart has a thicker wall, a far more diffused and thick distribution of tracheoles, and a far greater number of mitochondria with respect to the aorta. This arrangement ensures a greater availability of oxygen and energy in the abdominal heart compared to the aorta and leads one to suppose that the high‐ and low‐frequency contractions of the cardiac cycle (Thon, [1982] J. Insect Physiol. 28:411–416) can be attributed to the abdominal heart and the aorta, respectively. J. Morphol. 240:15–31, 1999. © 1999 Wiley‐Liss, Inc.     

Physical features of grass leaves influence the placement of eggs within the plant by the Hessian fly

Hessian fly eggs are more likely to be found on adaxial rather than abaxial surfaces of wheat leaves. These leaf surfaces differ in their physical features: the adaxial side of the leaf has parallel grooves and ridges while the abaxial side is relatively smooth. We used leaf models to investigate the relationship between Hessian fly egglaying and these physical features. When both sides of a green paper leaf model were treated with a wheat leaf extract, but only one side of the model was scored with parallel grooves, the grooved side received more eggs than the smooth side. As the number of grooves per surface increased from 0 to 10, eggs per model increased. When grooves and the wheat leaf extract were tested together and separately, the grooves significantly increased egg numbers in the presence, but not the absence, of wheat extract. In contrast, wheat extract increased egg numbers both in the absence and presence of grooves. Molding techniques were used to recreate the physical features of the adaxial and abaxial leaf surfaces of five grasses. For four of the grass genotypes (a triticale, two common wheats, and a durum wheat), patterns of egglaying on real leaves and molded models were similar, i.e., adaxial leaf surfaces and adaxial molded models were preferred over abaxial leaf surfaces and abaxial models. On the fifth grass, oat, preferences for the adaxial side of real leaves and for adaxial models were not as obvious. We conclude that the adult female Hessian fly obtains information about the leaf surface through her tactile and/or kinesthetic senses and uses this information when making egglaying decisions.     

Oviposition preferences of the Hessian fly and their consequences for the survival and reproductive potential of offspring

1. The relationship between oviposition preference and offspring performance was investigated experimentally for the Hessian fly Mayetiola destructor (Say) using 10 grass genotypes that represented five different genera and six different species. Oviposition preferences were quantified in a choice test using arrays of 200–400 plants. Offspring performance was estimated by recording survival during three phases of the insect–plant association and by measuring wing length, a correlate of adult reproductive potential. Density effects were examined for all offspring variables, and were taken into account when offspring performance was compared across the 10 grasses. 2. Egg counts from the oviposition choice test revealed a consistent ranking of plants: 18ITSN triticale > Otane bread wheat = Caldwell bread wheat = Stacy bread wheat > 3424 bread wheat = PND durum wheat = Fleet barley = Valetta barley = Matua brome grass > Awapuni oat. 3. Survival and adult wing length varied significantly on the 10 plant types. Survival was ranked: Otane bread wheat > 3424 bread wheat = Fleet barley = Matua brome grass > Caldwell bread wheat = Valetta barley > 18ITSN triticale = PND durum wheat = Stacy bread wheat > Awapuni oat. Adult male and female wing lengths were greatest on Otane, the bread wheat that also provided the best survival. 4. For Hessian flies on each of the 10 plant types, data on survival, wing length, and wing length–fecundity relationships were combined into a single fitness measure. When these fitness measures were compared with egg counts, no overall pattern emerged. On seven of the 10 plant types, there was a positive linear relationship between egg counts and offspring fitness. On the other three plant types, egg counts were high while fitness was low. Possible reasons why Hessian fly females lay eggs on plants that are poor hosts for their offspring are discussed.     

Characterization of two genes expressed in the salivary glands of the Hessian fly, Mayetiola destructor (Say)

Two genes, SSGP-11A1 and SSGP-12A1, have been isolated that encodes proteins with a secretion signal peptide at theN-terminals from the Hessian fly (Mayetiola destructor (Say)). The SSGP-11A1 gene contains one small intron (89 bp) and encodes a putative protein with 79 amino acids. The first 18 amino acids constitute a putative secretion signal peptide. The SSGP-12A1 gene contains three small introns and encodes a putative protein with 234 amino acids. The first 19 amino acids constitute a putative secretion signal peptide. Northern blot analysis revealed that both of the genes are primarily expressed in the salivary glands of Hessian fly larvae, the feeding stage of the insect. These observations are consistent with the possibility that the proteins encoded by them are secreted into host plants during feeding. Even though both genes are exclusively expressed in Hessian fly larvae, the expression profiles between them were quite different in insects at different instars. The SSGP-11A1 gene was expressed in all instars of larvae while the SSGP-12A1 gene was almost exclusively expressed in the first instar larvae. The differential expression suggests that the proteins encoded by these two genes may perform different functions. In situ hybridization revealed that the SSGP-11A1 gene is located on the short arm of chromosome A1 while SSGP-12A1 gene is on the long arm of chromosome A2.     

The larval head morphology of Xyela sp. (Xyelidae, Hymenoptera) and its phylogenetic implications

Larval head structures of Xyela sp. are described in detail. The characters are compared to conditions found in larvae of other groups of Hymenoptera and Endopterygota. Like other symphytan larvae the immature stages of Xyelidae are mainly characterized by presumably plesiomorphic features of the head. The head sutures are well developed and all parts of the tentorium are present. The labrum is free and a complete set of labral muscles is present. The maxillae are in a retracted position. In contrast to other hymenopteran larvae Xyela possesses a clypeofrontal suture, a comparatively long antenna and three well‐developed antennal muscles. Apomorphic features of Xyela are the absence of muscles associated with the salivarium and the complete absence of Musculus craniocardinalis. A clade comprising Orussidae and Apocrita is supported by the unsegmented maxillary and labial palps and the absence of the lacinia. Six potential autapomorphies for the Hymenoptera were revealed: (1) the caudal tentorial apodeme, (2) the bifurcated tendon of Musculus craniomandibularus internus, (3) the lateral lobe of the cardo, (4) the origin of M. tentoriohypopharyngalis from the posterior head capsule, (5) the exceptionally strong prepharyngo‐pharyngeal longitudinal muscle and (6) the longitudinal muscle of the silk press. The maxillolabial complex, the vestigial M. craniocardinalis and a distinctly developed labio‐hypopharyngeal lobe bearing the opening of the salivary duct are potential synapomorphies of Hymenoptera and Mecopterida. The globular, orthognathous head capsule, the modified compound eyes, the occipital furrow and the X‐shaped tentorium are features with unclear polarity shared by Hymenoptera and Mecoptera.     

The head morphology of Clambidae and its implications for the phylogeny of Scirtoidea (Coleoptera: Polyphaga)

External and internal structures of the head of adults of Clambus are described and illustrated in detail. The results are compared with structural features found in the clambid genus Calyptomerus, in representatives of other scirtoid families, and also in species of other coleopteran suborders, notably Myxophaga. The results tentatively support the monophyly of Scirtoidea and a close relationship between Clambidae and Eucinetidae is suggested by one shared derived feature of the mandible, a long and slender apical tooth with a serrate edge. The monophyly of Clambidae is very strongly supported and Acalyptomerus is probably the sistergroup of a clade Calyptomerus + Clambinae. Potential scirtoid autapomorphies are the loss of the dorsal tentorial arms, a bulging gula, a strongly transverse labrum, and a ridge separating the mediostipes from the lacinia. However, all these features are homoplasious. The monophyly of Clambidae is supported by modifications of the head capsule which is strongly flattened and broadened, by a deep clypeofrontal incision enabling vertical antennal movements, and a series of antennal features. Synapomorphies of Clambinae + Calyptomerus (Clambidae excluding Acalyptomerus) are the conglobate body form with the ventral side of the head capsule in contact with the mesocoxae, and compound eyes integrated in the contour of the head. The completely subdivided eye is an autapomorphy of Clambus. An entire series of features is shared by Clambidae (or Scirtoidea) and Myxophaga. Most of them are apomorphies that apparently evolved independently in both groups. However, the presence of well‐developed maxillary and labial glands is arguably a retained groundplan feature of Coleoptera, with parallel loss in Archostemata, Adephaga and various groups of Polyphaga. J. Morphol. 277:615–633, 2016. © 2016 Wiley Periodicals, Inc.     

Leaf growth signals the onset of effective plant resistance against Hessian fly larvae

For plant resistance that is induced rather than constitutive, the precise timing of a sequence of events must be considered (i.e., initial detection of the insect by the plant's surveillance systems, up-regulation of signaling and defense pathways, achievement of effective levels of defense, and finally down-regulation of signaling and defense). Here, we provide a timeline for the interaction between resistant wheat ( Triticum aestivum L.) (Poaceae) and the Hessian fly, Mayetiola destructor (Say) (Diptera: Cecidomyiidae). To create this timeline, we measured the daily growth of the third, fourth, and fifth leaves of susceptible and resistant plants. Because each leaf had a different spatial relationship to the site of larval attack (i.e., the sheath epidermal cells of the third leaf) and a different pattern of growth relative to the 3–5 days that larvae attacked resistant plants, we learned different things from each leaf. The third leaf shows how quickly responses of susceptible and resistant plants diverge (i.e., 36–60 h after initial larval attack). The fourth leaf shows that, for both susceptible and resistant plants, negative effects of larval attack extend beyond the third leaf. These negative effects are more severe for susceptible plants, but even in resistant plants continue for several days after larvae have died. The fifth leaf is interesting because it shows how rapidly the resistant plant recovers from larval attack. Thus, 204–348 h after initial attack, a time when the fourth leaf of resistant plants is showing reduced growth and the fifth leaf of susceptible plants is showing zero growth, the fifth leaf of resistant plants shows a small increase in growth. Grasses with resistance gene-mediated resistance may have a two-fold strategy, using resistance mechanisms to stop Hessian fly larvae from further attack and tolerance mechanisms to protect resources for future plant growth.     

突尼斯两种麦瘿蚊的遗传变异和相关性分析

小麦黑森瘿蚊Mayetiola destructor Say和大麦茎干瘿蚊M. hordei Kieffer是在突尼斯每年都可导致谷物重大损失的两个植食性姊妹种。通常认为为害小麦的瘿蚊是小麦黑森瘿蚊,但是不同谷类物种（小麦或大麦）与麦瘿蚊的这两个种（ destructor 或 hordei ）之间寄主关系并不很严格。提出有效的害虫管理方案首先要求对瘿蚊基因型进行精确分析。本研究应用随机扩增多态DNA（RAPD）技术,结合交配分析和线粒体DNA分型技术,对位于突尼斯北部的一个为害小麦的麦瘿蚊种群的遗传变异程度和分类关联性进行了评估。基于RAPD结果的系统发育分析表明,所研究的种群具有较大的遗传变异范围,这可能源于被分析的小麦样品有被两种瘿蚊共同侵害的复杂背景。虽然交配分析表明有少数不能成功产卵(2/14),但是基于细胞色素b基因限制性酶切分析显示全部样品的线粒体分型均属M. destructor。本文结果进一步证明以RAPD可变性作为分类推断依据不可靠,还为突尼斯M. destructor和M. hordei属于异域分布的观点提供了补充证据。     

Constructional morphology within the head of hammerhead sharks (sphyrnidae)

The study of functional trade‐offs is important if a structure, such as the cranium, serves multiple biological roles, and is, therefore, shaped by multiple selective pressures. The sphyrnid cephalofoil presents an excellent model for investigating potential trade‐offs among sensory, neural, and feeding structures. In this study, hammerhead shark species were chosen to represent differences in head form through phylogeny. A combination of surface‐based geometric morphometrics, computed tomography (CT) volumetric analysis, and phylogenetic analyses were utilized to investigate potential trade‐offs within the head. Hammerhead sharks display a diversity of cranial morphologies where the position of the eyes and nares vary among species, with only minor changes in shape, position, and volume of the feeding apparatus through phylogeny. The basal winghead shark, Eusphyra blochii, has small anteriorly positioned eyes. Through phylogeny, the relative size and position of the eyes change, such that derived species have larger, more medially positioned eyes. The lateral position of the external nares is highly variable, showing no phylogenetic trend. Mouth size and position are conserved, remaining relatively unchanged. Volumetric CT analyses reveal no trade‐offs between the feeding apparatus and the remaining cranial structures. The few trade‐offs were isolated to the nasal capsule volume's inverse correlation with braincase, chondrocranial, and total cephalofoil volume. Eye volume also decreased as cephalofoil width increased. These data indicate that despite considerable changes in head shape, much of the head is morphologically conserved through sphyrnid phylogeny, particularly the jaw cartilages and their associated feeding muscles, with shape change and morphological trade‐offs being primarily confined to the lateral wings of the cephalofoil and their associated sensory structures. J. Morphol. 276:526–539, 2015. © 2015 Wiley Periodicals, Inc.     

Genetic variation among populations of the Hessian fly Mayetiola destructor (Diptera: Cecidomyiidae) in Morocco and Syria

The RAPD-PCR technique was used to study genetic variation within and among geographical populations of the Hessian fly, Mayetiola destructor (Say), from Morocco and Syria, associated with the fly's ability to overcome resistance in three wheat cultivars containing H5, H13 and H22 resistance genes. Variation was detected both for the level of susceptibility of the cultivars and RAPD profiles of M. destructor populations. By the use of RAPD-PCR, high genetic variability was detected among individuals and populations of M. destructor within and between areas separated geographically. The DNA fingerprints of populations of M. destructor were area-specific with Nei's measures of genetic distance ranging from 0.156 (between Abda and Beni Mellal, Morocco) to 1.977 (between Marchouch, Morocco and Lattakia, Syria). Cluster analysis of the genetic distances among the populations, identified the Syrian population as an outlier. A highly significant correlation (r = 0.81) observed between the genetic and geographic distances among the populations, provided genetic support for dispersal of the fly from its presumed origin in West Asia to Morocco.     

The role of vision in the host plant-finding behaviour of the Hessian fly

Abstract. After mating, female Hessian flies, Mayetiola destructor (Say), sit for several hours before taking flight and exhibiting host-finding behaviours. Observation of these behaviours indicated that females used both chemical and visual cues to find oviposition sites, and that visual cues from plants consisted of both spectral (colour) and spatial information. Females oriented to and were stimulated to land on targets reflecting wavelengths in the region of 530-560nm (either alone or in combination with UV) but were inhibited by wavelengths from 400 to 500 nm. This spectral discrimination was probably mediated by photoreceptors located in the medio-frontal zone of the compound eyes. The spatial patterning of targets also influenced orientation during flight; targets with vertical rather than horizontal contour lengths and a higher density of vertical contour lengths were approached and landed on more frequently.     

中国伪安瘿蚊属一新种记述(双翅目,瘿蚊科)

记述了中国云南省的伪安瘿蚊属1新种:花椒伪安瘿蚊Pseudasphondylia zanthoxyli sp.nov.,模式标本存放于山东农业大学昆虫标本室.     

浙江端突瘿蚊属一新种记述(双翅目,瘿蚊科)

记述了采自浙江的端突瘿蚊属Epidiplosis 1新种,即长叉端突瘿蚊Epidiplosis furcata sp. nov..正模♂,浙江省西天目山,2000-07-19,墨铁路、刘涛灯诱.该新种与拟丝端突瘿蚊Epidiplosis simulifilifera Mo比较近似,与后者主要区别如下:1)抱器基节各具1叉状端瓣,阳具基侧突骨化,呈"T"形;2)下颚须第3节长约为宽的3.4倍;3)触角末鞭节的端结长为宽的2.5倍;4)体长1.16 mm.     

中国瘿蚊一新种记述(双翅目,瘿蚊科)

记述采自浙江省的端突瘿蚊属Epidiplosis 1新种:指状端突瘿蚊Epidiplosis dactylina sp.nov,.模式标本存放于山东农业大学昆虫标本室.     

Cloning and characterization of chymotrypsin- and trypsin-like cDNAs from the gut of the Hessian fly [Mayetiola destructor (Say)

Fifteen unique cDNA clones encoding trypsin- or chymotrypsin-like proteins were cloned and characterized from a gut cDNA library derived from Hessian fly [Mayetiola destructor (Say)] larvae. Based on sequence similarities, the cDNAs were sorted into five gene groups, which were named MDP1 to MDP5. Two of the gene groups, MDP1 and MDP2, encoded chymotrypsin-like proteins; the other three encoded putative trypsins. All deduced proteins have conserved His(87), Asp(136), and Ser(241) residues for the catalytic triad and three pairs of cysteine residues for disulfide bridge configurations. The substrate specificity determination residue at position 235 was also conserved in the putative trypsins and chymotrypsins. In addition, all the deduced protein precursors had a typical secretion signal peptide and activation peptide. Northern blot analysis revealed that all these gene groups were exclusively expressed in the larval stage. The expression profiles for each gene group differed significantly in different ages of the larva, as well as in different tissues. Protease activity analysis of gut extract, using specific inhibitors, demonstrated that serine proteases were the major digestive enzymes in the gut of M. destructor larvae. Serine protease inhibitors inhibited as much as 90% proteolytic activities of gut extract, whereas inhibitors specific to other proteases, including cysteine proteases, aspartic proteases, and metallo-proteases, inhibited only 10-24% of gut protease activity.