Selection for canalization at two extra dorsocentral bristles in Drosophila melanogaster.

From 10 isofemale lines of D. melanogaster, the D2 line was established with the aim of obtaining an invariant phenotype at two extra dorsocentral bristles. Line D2 was also subdivided into two other lines, SA and ASD, based on their different bristle patterns. The SA line was selected for two symmetrical anterior extra bristles, and the ASD line was selected for two asymmetrical extra bristles, one anterior and one posterior. Only the SA line showed any canalizing response (estimated by the width of the probit transformation) at the two-extra-bristle class. Nevertheless, the results from the different lines were more consistent with the independent ones of both the anterior and posterior regions of the extra dorsocentral bristles. This analysis showed some independent genetic systems for each region, developmental canalization being at two extra symmetrical bristles per region in all the selected lines (D2, ASD, and SA). Therefore, this canalization did not depend directly on the extra-bristle positional pattern used in the selection. The wild-type canalizing system is suggested to explain the fast canalizing response in a phenotype that had not been previously canalized by natural selection.     

Selection for canalization at extra dorsocentral and scutellar bristles in Drosophila melanogaster.

Direct artificial selection for a specific pattern, in the number and position of extra bristles, was carried out in a wild-type population of Drosophila melanogaster to canalize (estimated by probit width) the selected phenotypes. From the same population, independent lines were selected for extra dorsocentral bristles (lines D3 and D4) and for extra scutellar bristles (lines E2, E3, and E4). Differences at canalization between both dorsocentral and scutellar systems were detected. Results fit an independent control hypothesis for canalization, at two symmetrical extra bristles, in the main regions in which extra bristles appear.     

Selection for a Threshold Character in Drosophila IV. Chromosomal Analyses of Plateaued Populations

Chromosome substitution and intra-chromosomal recombination techniques have been employed to determine the genetic basis of limits to selection in lines selected for high scutellar bristle number from the Canberra population. Three observations indicate the presence of an upper threshold affecting some component traits, which is not readily discernible at the level of the selected phenotype:(1) The variance of the number of anterior + interstitial + posterior bristles is progressively reduced as the mean approaches a total of eight at these sites. Total bristle number, which includes apicals in addition to the above three components, and which was the trait subject to selection, shows little evidence of this phenomenon;(2) the effect of a given chromosome substitution is also greatly reduced as the mean approaches eight anterior + interstitial + posterior bristles, by comparison with its effect in genotypes of lower mean;(3) chromosome substitutions show some evidence of negative interaction as this level is approached, in contrast to the positive interactions evident at higher means. All chromosomes except IV are involved in progress beyond the proposed upper threshold. However, chromosome III has the most important effect, due primarily to a major gene located at approximately 61 cM, which also markedly increases dorsocentral and postvertical bristle numbers.     

Chromosomal polymorphism and extra bristles of Drosophila melanogaster: joint variation under selection in isofemale lines

Selection for increased numbers of dorsocentral and scutellar bristles produced an increase of In(3R)C heterozygotes in isofemale lines of D. melanogaster. The influence of polygenic selection in the maintenance of the chromosomal polymorphism is discussed.     

Canalization at two extra scutellar bristles: artificial selection, chromosome analysis, and effect of temperature.

Two isofemale lines (P1 and P2) from a wild-type population of Drosophila melanogaster were selected for an invariant phenotype, two symmetrical and anterior scutellar extra bristles. Both P1 and P2 lines responded quickly to direct selection for two extra bristles, but although this phenotype was canalized in the P2 line, it was not in the P1 line. This lack of canalization at two extra bristles, measured by probit width, is the first reported in the literature. Analysis of chromosomal contribution showed the P1 decanalized phenotype to be due to chromosome 2. Synthetic chromosomal combinations were tested at four development temperatures (17 degrees, 21 degrees, 24 degrees, and 30 degrees C), and no correlation was observed between canalization at two extra bristles measured by probit width and minimum phenotypic change due to different temperatures. In this test, development at the highest temperatures was accompanied by an increased tendency to return to the wild phenotype in the canalized phenotypes only, suggesting that this drastic change could be accounted for by the action of the wild-type canalizing system. Decanalized genotypes, or those canalizing at phenotypes other than the wild type, could be explained by mutations which, to a greater or lesser degree, alter the normal genetic system of canalization.     

STABILIZING SELECTION DETECTED FOR BRISTLE NUMBER IN DROSOPHILA MELANOGASTER

Stabilizing selection, which favors intermediate phenotypes, is frequently invoked as the selective force maintaining a population's status quo. Two main alternative reasons for stabilizing selection on a quantitative trait are possible: (1) intermediate trait values can be favored through the causal effect of the trait on fitness (direct stabilizing selection); or (2) through a pleiotropic, deleterious side effect on fitness of mutants affecting the trait (apparent stabilizing selection). Up to now, these alternatives have never been experimentally disentangled. Here we measure fitness as a function of the number of abdominal bristles within four Drosophila melanogaster lines, one with high, one with low, and two with intermediate average bristle number. The four were inbred nonsegregating lines, so that apparent selection due to pleiotropy is not possible. Individual fitness significantly increased (decreased) with bristles number in the low (high) line. No significant fitness-trait association was detected within each intermediate line. These results reveal substantial direct stabilizing selection on the trait.     

Selection and temperature effects on extra dorsocentral bristles inDrosophila melanogaster

Starting from base populations which showed a tendency to form supernumerary dorsocentral bristles, selection for high numbers of dorsocentrals was carried out. In all, 9 lines were selected 6 to 14 generations. Selection proved to be effective in all but two lines. Selection for reduced numbers in one of the lines also proved effective.Mean values of over 10 were reached in the females of certain high lines. Average counts in males were always lower than those of females. This sex difference is not a consequence of the difference in size between the sexes.The base populations were of mutant stock origin, and some of them segregated for mutant genes which proved to be correlated with extrabristle phenotype. Supernumerary bristles were not distributed in the same manner in all of the selection lines, nor was the reaction of phenotypic expression to temperature the same in various lines.     

A new genus and a new species of tachinids (Diptera,Tachinidae) from the south of middle Asia

Ventoplagia gen. n. is described, with the type species Ventoplagia brevirostris sp. n. The frontal bristles extending only to the base of the pedicel, 2+3 dorsocentral bristles, 0+2 intraalar bristles, the absence of prealar bristle, the short and fine anepimeral (pteropleural) bristle, the scutellum without lateral bristles, and the welldeveloped posteroventral bristle of the hind tibia indicate that the new genus belongs to tribe Minthoini. Ventoplagia gen. n. is closely related to the genus Palmonia Kugler. The characters distinguishing these genera are given.     

Shaggy (zeste-white 3) and the formation of supernumerary bristle precursors in the developing wing blade of Drosophila.

The development of supernumerary bristle precursors induced by the mutation shaggy (sgg; also known as zeste-white 3) was examined in the developing wing blade of imaginal and pupal Drosophila. sgg clones were induced by mitotic recombination; clones were marked using enhancer-trap flies which express beta-galactosidase ubiquitously in imaginal tissues, while bristle precursors were identified using sensillum and bristle-specific enhancer-trap lines. It was shown that the precursors of supernumerary sgg bristles in the wing blade mimicked the development of morphologically similar margin bristles, developing in a manner similar to that of anterior sensory bristles in anterior clones and posterior noninnervated bristles in posterior clones. Interestingly, supernumerary anterior sensory bristles appeared outside the normal regions of "proneural" gene activity as identified using anti-achaete. Moreover, sgg could induce the ectopic expression of achaete in anterior clones. Thus, in the anterior wing blade the sgg mutation leads to the formation of ectopic proneural regions.     

SECRETION AND DEPLOYMENT OF BRISTLES IN MALLOMONAS SPLENDENS (SYNUROPHYCEAE)1

Mallomonas splendens (G. S. West) Playfair has a cell covering of siliceous scales and bristles. Interphase cells bear four anterior and four posterior bristles that each articulate, at their flexed basal ends via a complex of labile fibers (the fibrillar complex), on a specialized body scale (a base-plate scale). Body scales, base-plate scales and bristles are formed independently of each other and at different times in silica deposition vesicles (SDVs) that are associated with one of the two chloroplasts. The fine structure of scale and bristle morphogenesis in M. splendens agrees with that previously described for Synura and Mallomonas. Four new posterior bristles are formed at late interphase with their basal ends towards the cell posterior. The fibrillar complex is formed in situ on the bristle in the SDV. Mature bristles are secreted one by one onto the surface of the protoplast, beneath the layer of body scales, where the basal ends of the bristles adhere to the plasma membrane via the fibrillar complex. The extrusion of posterior bristles and their deployment onto the cell surface was monitored with video. A fine cellular protuberance accompanies the bristles as they are extruded from beneath the scale layer with their basal ends leading. When distant from the cell, the basal ends of the bristles appear attached to the protuberance, possibly by way of their fibrillar complexes. Once bristles are fully extruded, and their tips free in the surrounding environment, the bristle bases are drawn back to the posterior apex of the cell, apparently by the now shortening protuberance. Thus a 180° reorientation of the posterior bristles has been effected outside the cell. Thin-sections of cells that are extruding bristles show a threadlike, cytoplasmic extension of the cell posterior which may be analogous to the protuberance seen in live cells. Four new posterior base-plate scales are secreted after the bristles have reoriented. Scanning electron microscopy indicates that the fibrillar complex is involved in positioning the bristles onto their respective base-plate scales. Anterior bristles are formed in new daughter cells in the same orientation as the posterior bristles; thus they are extruded tip first and no reorientation is required.     

Analysis of lethals in selected lines of Drosophila melanogaster

Summary Five lines of Drosophila melanogaster that reached an extreme phenotype after long-term selection for increased dorsocentral bristle number, were analysed for the presence of lethals. Seven chromosome II and three chromosome III lethal types were detected in four of the lines, at frequencies ranging from between 6% and 36%. No lethal had any demonstrable effect over the selected trait. In one line, where almost every chromosome II was a lethal carrier, it was shown that the main lethal (at a frequency of 36%) was associated with the transmission ratio distortion in males. The processes which could lead to the accumulation of this lethal and others linked in disequilibrium to it is discussed. Some results suggest similar mechanisms for the accumulation of lethals in the other lines. These findings show that causes other than the direct effect of artificial selection must be taken into account when trying to explain the accumulation of lethals in selected lines.     

Mesosternal bristle number in a cosmopolitan drosophilid: an X-linked variable trait independent of sternopleural bristles

Mesosternal (MS) bristles in Drosophila are a pair of machrochaetae found at the sternal end of the sternopleural (STP) microchaetae, and are thought to be invariable. In a closely related drosophilid genus, Zaprionus, their number is four and, in contrast to Drosophila, they show interspecific and intraspecific variability. The genetic basis of MS bristle number variability was studied in Z. indianus, the only cosmopolitan species of the genus. The trait responded rapidly to selection and two lines were obtained, one lacking any bristles (0-0) and the other bearing the normal phenotype (2-2). Other symmetrical phenotypes, (1-1) and (3-3), could also be selected for, but with lesser success. By contrast, STP bristle number did not vary significantly between the two lines (0-0) and (2-2), revealing its genetic independence from MS bristle number. Reciprocal crosses between these two lines showed that MS bristle number is mainly influenced by a major gene on the X chromosome (i.e. F₁ males always resembled their mothers) with codominant expression (i.e. heterozygous F₁ females harboured an average phenotype of 2 bristles). However, trait penetrance was incomplete and backcrosses revealed that this variability was partly due to genetic modifiers, most likely autosomal. The canalization of MS bristle number was investigated under different temperatures, and the increased appearance of abnormal phenotypes mainly occurred at extreme temperatures. There was a bias, however, towards bristle loss, as shown by a liability (developmental map) analysis. Finally, when ancestral and introduced populations were compared, the latter were far less stable, suggesting that genetic bottlenecks may perturb the MS bristle number canalization system. MS bristle number, thus, appears to be an excellent model for investigating developmental canalization at both the quantitative and the molecular level.     

Studies on the Scutellar Bristles of DROSOPHILA MELANOGASTER. II. Long-Term Selection for High Bristle Number in the Oregon Rc Strain and Correlated Responses in Abdominal Chaetae

Results are presented of 135 generations of selection for high scutellar bristle number in two lines M and M3 derived from the same original mating of one female with 5 bristles by one male with 4 bristles, the latter being the wild-type canalised phenotype. Results are also given of two relaxed lines per line and of a reselection line M2 derived from the first relaxed line of line M which had regressed almost to base population level. The effect of introducing the sc(1) allele into the M and M3 selected backgrounds was studied at generations 39-44. At the end of selection the effect of an extra dose of sc(+) was also studied in males of all selected backgrounds. The correlated responses in abdominal bristles were followed in all lines.-Considering their common origin, the selection lines differed markedly in pattern of scutellar response and in most other aspects observed, namely correlated responses in abdominals and p.c. scutellars, sex differences, and behaviour on relaxation. Selection limits for scutellar bristles in lines M and M2 were equal to or greater than the most extreme reported in the literature.-The probit span of the canalised 4 bristle class decreased in each selection line as the mean scutellar bristle number increased, and increased again in the relaxed lines as the mean bristle number decreased. In the context of an hypothesis that canalisation at 4 bristle is due to regulation of the scute locus, this result is now interpreted as being due mainly to selection for poor regulators of sc(+), in contrast to a previous interpretation that only the minor gene background was altered by selection, the canalisation (regulation) genotype not being affected.-Introducing the sc(1) allele into the selected backgrounds M and M3 showed a reduced effect on sc(1) flies compared with sc(+) flies, and an interaction of sc(1) and sc(+) with selected background. sc(1) flies had about the same number of bristles in both backgrounds though the mean of sc(+) flies in line M was about 3sigma higher than in line M3. Dominance of sc(+) to sc(1) was reduced slightly in M3. However, the effect of an extra dose of sc(+) at the end of selection was about the same as in unselected in all lines, so the first or dominance level of regulation of the scute locus was not significantly affected by selection, though the second or canalisation level of regulation was.-A large positive correlated response in abdominal bristles occurred in all lines. The response in line M was about twice that in M2 and M3 and was in fact as large as can be obtained from direct selection on abdominals. In line M some genes may have been selected with a proportionately greater effect on abdominals than on scutellars. This is supported by the further observation in line M that the abdominal scores of flies with particular scutellar bristles scores increased as the scutellar mean increased. An attempt was made to apply to these results Rendel's (1962) model of competition between scutellars and abdominals for common bristle-making resources. This could not be done satisfactorily mainly because the assumptions in the model about the similarity of effects in scute and wild-type flies were not met in the present material.     

Selection on bristle length has the ability to drive the evolution of male abdominal appendages in the sepsid fly Themira biloba

Many exaggerated and novel traits are strongly influenced by sexual selection. Although sexual selection is a powerful evolutionary force, underlying genetic interactions can constrain evolutionary outcomes. The relative strength of selection vs. constraint has been a matter of debate for the evolution of male abdominal appendages in sepsid flies. These abdominal appendages are involved in courtship and mating, but their function has not been directly tested. We performed mate choice experiments to determine whether sexual selection acts on abdominal appendages in the sepsid Themira biloba. We tested whether appendage bristle length influenced successful insemination by surgically trimming the bristles. Females paired with males that had shortened bristles laid only unfertilized eggs, indicating that long bristles are necessary for successful insemination. We also tested whether the evolution of bristle length was constrained by phenotypic correlations with other traits. Analyses of phenotypic covariation indicated that bristle length was highly correlated with other abdominal appendage traits, but was not correlated with abdominal sternite size. Thus, abdominal appendages are not exaggerated traits like many sexual ornaments, but vary independently from body size. At the same time, strong correlations between bristle length and appendage length suggest that selection on bristle length is likely to result in a correlated increase in appendage length. Bristle length is under sexual selection in T. biloba and has the potential to evolve independently from abdomen size.     

Effects of different base populations on selection response in Drosophila

Of two laboratory strains of Drosophila melanogaster used in this study, the +3 strain had slightly higher mean abdominal bristle number and estimated heritability of this character than the Oregon‐R. Their F₁ hybrid exhibited 5 % heterosis. Fourteen generations of the two original strains and the F₃ of the hybrid were selected for high and low numbers of abdominal bristles on the 4th and 5th sternites, at a selection intensity of 20%. A mass‐mated unselected control was maintained for each population. The +3 population responded considerably more to selection for low numbers of bristles than high, and the Oregon‐R population showed a similar, though less marked, tendency; the Crossbred population responded more strongly to selection for high numbers. Except for the Crossbred high selection line, all lines declined in response rate, phenotypic variance, and realised heritability. The average realised heritability of the Oregon‐R and +3 high and low selection lines over 14 selection generations fell short of their predicted base population heritabilities. The deviation from the predicted was particularly pronounced with selection for high bristle number in the +3 line.     

Updating the Trachelocercids (Ciliophora, Karyorelictea). I. A Detailed Description of the Infraciliature of Trachelolophos gigas N. G., N. Sp. and T. filum (Dragesco & Dragesco-Kernéis, 1986) N. Comb

ABSTRACT. Trachelolophos gigas n. g., n. sp. and T. filum (Dragesco & Dragesco-Kernéis, 1986) n. comb. (basionym: Tracheloraphis filum) were discovered in the mesopsammon of the French Atlantic coast at Roscoff. Their morphology and infraciliature were studied in live and protargol impregnated specimens. The new genus, Trachelolophos, belongs to the family Trachelocercidae and is unique in having a conspicuous ciliary tuft, which is very likely a highly modified brosse, in the oral cavity. The two species investigated have a very similar infraciliature, differing only in morphometric characteristics and in the nuclear configuration. The entire somatic and oral infraciliature consists of dikinetids which have both basal bodies ciliated or only the anterior or posterior ones, depending on the region of the cell. The right side is densely and uniformly ciliated. Its kineties extend onto the left side to the glabrous stripe, where an anterior and posterior secant system are formed, reducing the number of kineties in the narrowed neck and tail region. The left side bears a narrow glabrous stripe bordered by slightly irregularly arranged dikinetids having rather stiff cilia (bristles), possibly forming an uninterrupted, prolate ellipsoidal (bristle) kinety as indicated by their ciliation. The bristle kinety commences subapically at the right margin of the glabrous stripe, extends posteriorly, then anteriorly at the left, to end up at the right margin again. The dikinetids of the right posterior portion of the bristle kinety have the posterior basal bodies ciliated, whereas the anterior basal bodies are ciliated in its left and right anterior portion. The ends of the bristle kinety meet distinctly subapically at the right margin of the glabrous stripe, as indicated by the diametrically opposed ciliation of the dikinetids. The anterior region (head) of the cell bears a distinct circumoral kinety composed of very regularly arranged dikinetids, associated with nematodesmata forming an oral basket together with the nematodesmal bundles originating from the oralized somatic dikinetids at the anterior end of the somatic kineties. The systematics of trachelocercid ciliates are briefly reviewed and discussed.     

Ontogeny of the complex sperm in the macrostomid flatworm Macrostomum lignano (Macrostomorpha,Rhabditophora)

Spermiogenesis in Macrostomum lignano (Macrostomorpha, Rhabditophora) is described using light‐ and electron microscopy of the successive stages in sperm development. Ovoid spermatids develop to highly complex, elongated sperm possessing an undulating distal (anterior) process (or “feeler”), bristles, and a proximal (posterior) brush. In particular, we present a detailed account of the morphology and ontogeny of the bristles, describing for the first time the formation of a highly specialized bristle complex consisting of several parts. This complex is ultimately reduced when sperm are mature. The implications of the development of this bristle complex on both sperm maturation and the evolution and function of the bristles are discussed. The assumed homology between bristles and flagellae questioned. J. Morphol., 2009. © 2008 Wiley‐Liss, Inc.