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Biomedical subjects

A Hoikkala

Publications and source records attributed to A Hoikkala.

At least 19 recordsLinked to original sources

Enterolignans.

A review with 114 references about mammalian lignans (enterolignans). Several aspects have been reviewed: the precursors of mammalian lignans and their biosynthesis, biological activities and health effects, metabolism (in vivo and in vitro) in human and animals, some synthetic strategies to obtain enterolignan skeleton types, including the synthesis of haptens and deuterated lignans, and finally an overview of the analytical methods to detect and quantify lignans in biological matrices and foods.

Animals↗

Nucleotide variation at the no-on-transient A gene in Drosophila littoralis.

The no-on-transient A (nonA) gene encodes a putative RNA-binding protein, and mutations in this gene are known to affect vision, male courtship song and viability in Drosophila melanogaster. Here we have sequenced the coding region of the nonA gene of Drosophila littoralis and compared it with those of Drosophila virilis and D. melanogaster. All portions of nonA appeared to be conserved between D. littoralis and D. virilis, while the 5' region of the gene of these two species showed high divergence from that of a more distantly-related species, D. melanogaster. The same was true for the glycine repeat regions. No significant deviation from neutrality was observed in the analysis of intraspecific nucleotide variation in 5' or 3' region of the nonA gene in D. littoralis population. Also, comparison of D. littoralis sequences with homologous sequence of D. virilis suggests that the gene is evolving neutrally in D. virilis group. Divergence of the 5' regions between D. virilis group species and D. melanogaster could be a result of positive selection, but this finding is obscured by the long divergence time of the species groups.

Animals↗

Variability levels, population size and structure of American and European Drosophila montana populations.

The level and patterns of nucleotide diversity have been characterized for two X-linked loci, fused (fu; a region of 2362 bp) and suppressor of sable (su(s); a region of 413 bp), in one European and one American D. montana population. Sequence variation at these loci shows that the two populations are divergent, although they may not be completely isolated. Data on the level of silent site variability at su(s) (1.1% and 0.5% for the European and American populations, respectively) suggest that the effective population sizes of the two populations may be similar. At the fused locus, one European sequence was highly divergent and may have resulted from gene conversion, and was excluded from the analysis. With this sequence removed, the level of silent site variability was significantly lower in the European population (0.28%) than in the American population (2.3%), which suggests a selective sweep at or near fu in the former population.

Animals↗

Identification of X chromosomal restriction fragment length polymorphism markers and their use in a gene localization study in Drosophila virilis and D. littoralis.

We have identified six restriction fragment length polymorphism (RFLP) markers based on unique gene sequences on the X chromosome of Drosophila virilis and D. littoralis. The markers were localized by in situ hybridization on larval polytene chromosomes, and the conjugation of the X chromosomes of the two species was studied in salivary glands of interspecific hybrid female larvae. The gene arrangement of D. virilis and D. littoralis appeared to be very different at the proximal end of the X chromosome preventing recombination between RFLP markers located in this area. Simple quantitative trait loci (QTL) analysis showed that five of our marker genes (including nonA and Dmca1A, previously found to affect male courtship song in D. melanogaster) are linked with a gene(s) having a major effect on species differences in the male courtship song between D. virilis and D. littoralis. This shows that the song gene(s) may be located inside a large X-chromosomal inversion in D. littoralis (as previously suggested), but that it may also be located on an area between this inversion and the centromere, close to nonA and Dmca1A. Localization of this gene or gene complex will be continued with the aid of our newly identified RFLP markers by making interspecific crosses between D. virilis group species with more similar X chromosomes.

Animals↗

Characterization of female preference functions for Drosophila montana courtship song and a test of the temperature coupling hypothesis.

Female mate preferences are a major cause of diversity and elaboration in male sexual traits. Here we characterize the shape of female preference functions for pulse length and carrier frequency of the courtship song of Drosophila montana by fitting both parametric and nonparametric functions to the incidence of female receptive gestures to synthetic song. Preference functions for both traits are strongly directional. That for pulse length is linear and favors short pulses, whereas that for carrier frequency is stabilizing in shape, but would exert directional preferences favoring males with high carrier frequency. The preference for carrier frequency has probably evolved under sexual selection, but reasons for the preference for short pulses are less apparent. We also examine the effect of ambient temperature on the carrier frequency of male song and on the preference function for carrier frequency. For many similar acoustic communication systems, temperature coupling, a compensatory effect of temperature on preference functions, is thought to maintain coordination between preferences and signals. However, although the carrier frequency of D. montana song is highly dependent on environmental temperature, there is no temperature coupling of the female preference function. We suggest that temperature coupling may often arise due to a common effect of temperature on song and preference, rather than be an advantageous characteristic whose function is to maintain coordination in temperature-affected communication systems.

Animals↗

SEM search for sound production and sound perception organs in a variety of Drosophila species.

The discovery of new song types in Hawaiian Drosophila species has raised a question of alternative sound production and sound perception mechanisms in Drosophila. For example D. disjuncta males, which produce song with a carrier frequency of 5,000-6,000 Hz, do not vibrate their wings but only tremble them in a very small amplitude while singing. In our SEM search we found at the wing base of the flies of this species a special structure which could play a part in song production. Our search for auditory sense organs in Drosophila species revealed that the aristae on fly antennae are structurally not as simple as has been assumed. In Hawaiian D. planitibia subgroup species, the fly aristae are more regular than in other Drosophila species and they are also covered with small hairs. We did not find any sign of tympanic hearing organs in any of the studied species, but we discovered on fly legs and wings sensilla which could play a part in sound/vibration perception. Our SEM survey provides a good starting point in the search for alternative pathways for sound production and perception via transmission electron microscopy combined with behavioral studies and electrophysiological recordings in Hawaiian Drosophila species.

Animals↗

Localization of genes affecting species differences in male courtship song between Drosophila virilis and D. littoralis.

The males of six species of the Drosophila virilis group (including D. virilis) keep their wings extended while producing a train of sound pulses, where the pulses follow each other without any pause. The males of the remaining five species of the group produce only one sound pulse during each wing extension/vibration, which results in species-specific songs with long pauses (in D. littoralis about 300 ms) between successive sound pulses. Genetic analyses of the differences between the songs of D. virilis and D. littoralis showed that species-specific song traits are affected by genes on the X chromosome, and for the length of pause, also by genes on chromosomes 3 and 4. The X chromosomal genes having a major impact on pulse and pause length were tightly linked with white, apricot and notched marker genes located at the proximal third of the chromosome. A large inversion in D. littoralis, marked by notched, prevents more precise localization of these genes by classical crossing methods.

Animals↗

Variation and consistency of female preferences for simulated courtship songs in Drosophila virilis.

To study the variation and consistency of song preferences in Drosophila virilis females, we played them species-specific song and songs with modified sound pulses and/or interpulse intervals on 3 consecutive days. Species-specific song was played again on the fourth day. All playbacks were done without the presence of males. About 62% of the females indicated their readiness to mate by spreading their wings in at least one of the trials. The proportion of the females responding to species-specific song was about twice that of the females responding to modified songs. The majority of females responded to only one song type, which suggests that the females varied in their preferences and that their preference windows were rather narrow. The females were consistent in their responsiveness to species-specific song played on 2 days. If the female responded to normal song during the first trial, the probability of her responding to the same song during the second trial increased by about 32%. The number of songs required by the females before responding in the two subsequent trials was also correlated within the females (repeatability 0.328). Repeatability of female preferences for male sexual traits is expected both in the viability and Fisherian models of sexual selection. Copyright 1999 The Association for the Study of Animal Behaviour.

Journal Article↗

Variability and evolvability of male song characters in Drosophila montana populations.

In Drosophila montana, the male courtship song (especially the pulse characters of the song) plays an important role in sexual selection. The heritabilities and the amount of additive and residual variation in different characters of the male song were measured in two populations using father-son regression and sib analysis. The songs of the males from the Oulanka population were recorded for a second time after the males had been kept in 4 degrees C for 6 months. Heritabilities measured for different song traits were non-significant in each case, largely due to high residual variation. During the cold treatment, the additive variation increased and the residual variation decreased in nearly all song traits. Our results suggest that genotype-environment interactions may increase the amount of additive variation between males in sexually selected song traits during overwintering, prior to the mating season of the flies.

Animals↗

Inheritance of species differences in female receptivity and song requirement between Drosophila virilis and D. montana.

Females of two Drosophila virilis group species, D. virilis and D. montana, have different requirements for the courting males. In the present study we have examined species differences in female receptivity and male courtship song requirement using females' acceptance signal instead of copulation for measuring female readiness to mate. Behavior of D. virilis and D. montana females and F1 and backcross hybrid females was observed in a single-pair courtships with D. virilis and D. montana males and normal and wingless (mute) F1 hybrid males. D. virilis females were very receptive and they commonly accepted the courtship of males unable to produce courtship song. D. montana females, on the contrary, had a low receptivity and these females accepted the courting male only after hearing his song. Interspecific F1 and backcross (BCm) females resembled D. virilis more than D. montana in their receptivity. These females, however, resembled D. montana in their song requirement. These findings suggest that female song requirement and female receptivity are determined by different genetic factors.

Animals↗

Male courtship song frequency as an indicator of male genetic quality in an insect species, Drosophila montana.

Most theoretical models on evolution of male secondary sexual characters and female preferences for these characters suggest that the male characters evolve in response to female preferences that may themselves evolve in response to direct or indirect benefits of choice. In Drosophila montana (a species of the D. virilis group), females use male song in their mate choice, preferring males that produce songs with short sound pulses and a high carrier frequency. We demonstrate here that the females get indirect benefits from their choice: in our data the frequency of the male song correlated with the survival rate of the male's progeny from egg to adulthood (indirect benefit for the female), but not with the fecundity of his mating partner (no direct benefit for the female). Male wing centroid asymmetry did not correlate with male wing song characters, nor with female egg production nor the fitness of her progeny, suggesting that fluctuating asymmetry in male wings does not play a major role in sexual signalling. The fact that the male song gives the female information on the male's condition/genetic quality in D. montana suggests that in this species the evolution of female preferences for male song characters could have evolved through condition-dependent viability selection presented in some 'good genes' models.

Animals↗

Female preference for fly song: playback experiments confirm the targets of sexual selection.

The courtship song of Drosophila is thought to be involved in sexual selection and species recognition. Because of the mating system of flies, however, directly demonstrating that song influences female preference is difficult. The majority of previous studies have used an experimental design that potentially confounds male and female reactions to song. In D. montana, correlational evidence has suggested that males that produce short sound pulses consisting of a high number of sound cycles (i.e. a high carrier frequency) have a higher mating success than other males. In this study, we played synthetic song that varied in pulse length and carrier frequency to individual females in the laboratory, both alone and in the presence of mute males. We scored female preference via an acceptance posture, 'wing spreading', which the females of this species usually display prior to mounting by a male. Females responded to synthetic song in the absence of males. The presence of mute males significantly increased their overall responsiveness, but the relative effectiveness of the songs did not change, eliminating male reaction to song as a possible confounding factor in the results. The interaction between pulse length and carrier frequency determined the discrimination between song types, with females responding most readily to song consisting of short pulses with a high carrier frequency. Thus, direct examination of female preferences supports the previous studies of male mating success, and confirms female song preference as a likely determinant of male mating success. Copyright 1998 The Association for the Study of Animal Behaviour

Journal Article↗

Songs produced by the females of the Drosophila virilis group of species.

Females of the Drosophila virilis group of species may vibrate their wings during courtship producing songs consisting of trains of successive sound pulses (pulse song) or sinusoidal hums (sine song). In the present study we describe these songs and study their role in courtship using a transition analysis. To find out which songs should be classified as pulse songs and which ones as sine songs, we studied the inheritance of different song types in interspecific F1 hybrids. In only a few species did the females produce large quantities of song in successful courtships. The males' reactions to female songs varied from licking and singing to stopping the courtship. Pulse trains with short and long intervals between successive pulses appeared to be different forms of the pulse song, while sine song (sinusoidal hums) was inherited independently of the pulse song.

Animals↗

Secondary courtship songs and inhibitory songs of Drosophila virilis-group males.

The males of all Drosophila virilis-group species produce primary courtship song; and the males of four of these species also provide secondary courtship song when courting a female. The amount of secondary song and the courtship phase at which it is produced vary according to the species. D. lummei males produce secondary song consisting of successive 12-ms-long sound pulses with 70-ms-long intervals between pulses. D. borealis males produce short and dense pulse trains and D. littoralis and D. flavomontana males produce single sound pulses of a long duration (80-160 ms). The males of all species produce pulse-structured inhibitory song when they are courted by another male. In the secondary and inhibitory songs of interspecific hybrids, short and dense pulse trains of parent species break into pulse song with long interpulse intervals. This suggests that songs consisting of short and dense pulse trains and pulse songs with long interpulse intervals are just different modes of the same song. Sine songs (long sound pulses) seem to be inherited independently of pulse song.

Animals↗

Change in the signal-response sequence responsible for asymmetric isolation between Drosophila planitibia and Drosophila silvestris.

Drosophila planitibia and Drosophila silvestris form a species pair that is an example of species diverged through a founder event. These species exhibit asymmetric sexual isolation, courtships between D. planitibia males and D. silvestris females being more successful than courtships between D. silvestris males and D. planitibia females. When analyzing the signal-response courtship sequence in these species, we found that D. silvestris females responded to male circling by standing or preening while D. planitibia females required further signals from the male to stop walking. The main reason for the reduced mating success rate of D. silvestris males with D. planitibia females was that the females responded to male circling by walking and the males did not proceed to the head-under-wings (HUW) position of a walking female. Another critical phase in these courtships was the HUW position in D. silvestris, where males proceeded almost immediately to wing and leg vibration. The courtships between D. planitibia male and D. silvestris female proceeded in a signal-response coordination until the male went to the HUW position, where he fanned his wings for too long a period before proceeding to wing and leg vibration. Thus, it seems that the asymmetric isolation between D. planitibia (ancestral species) and D. silvestris (derived species) is mainly due to a loss of transitions in the signal-response chain of D. silvestris. A change in the behavior of the males in the HUW position has caused further isolation between the species in both directions.

Animals↗

Laboratory and natural heritabilities of male courtship song characters in Drosophila montana and D. littoralis.

We estimated heritabilities for several male courtship song characters in two Drosophila species using father-son regression under conditions where both fathers and sons had been raised in the laboratory. In D. montana the heritabilities of song characters were rather high (-0.23 to 0.80) and in most cases significant. In D. littoralis the heritabilities of song characters were generally lower (-0.33 to 0.18), and none of them was significantly larger than zero. We also estimated heritabilities regressing characters of wild-caught fathers with those of their laboratory reared sons, and used the method employed by Riska et al. to estimate the lower bound of heritabilities in nature. In D. montana most and in D. littoralis all of the across-environment heritabilities were non-significant (-0.15 to 0.43 and -0.04 to 0.15, respectively), and in some cases the across-environment heritabilities were significantly lower than the heritabilities measured under laboratory conditions. The low across-environment heritabilities appeared to be due to larger phenotypic variability of song characters in the field and in some cases also due to genotype-environment interactions.

Animals↗

Inheritance of male courtship sound characteristics in Drosophila littoralis.

Males of Drosophila littoralis vibrate their wings during courtship to deliver a "love song." This consists of 25- to 50-ms-long pulses with a basic frequency of about 250-400 Hz, separated by 250- to 500-ms pauses. When recording the sounds of flies from several localities in Europe, we found that males of one strain from northern Finland displayed courtship sounds with an unusually low wing beat frequency (below 250 Hz). In a genetic analysis utilizing marker stocks, the anomalous frequency was found to be caused by genes on all major autosomes, the strongest factors being on the second chromosome. Interaction between genes on chromosome 2 and on the fused chromosome 3-4 was non-additive. In low-frequency sounds, the number of cycles in the pulse (CN) was decreased, so that the length of the sound pulse (PL) remained more or less unchanged. We suggest that the genetically and physiologically most thoroughly controlled trait in the sound of Drosophila littoralis is the length of the pulse.

Animals↗

Hawaiian courtship songs: evolutionary innovation in communication signals of Drosophila.

In Hawaii, flies of the genus Drosophila have undergone spectacular adaptive radiation, resulting in the evolution of more than 500 species of Drosophila that are found nowhere else on earth. This taxonomic uniqueness is reflected in behavior and morphology. Hawaiian Drosophila sing songs, as do continental Drosophila; however, the Hawaiian songs have diverged strongly in form and mechanism of production. The click-song of D. fasciculisetae's (Maui) has a carrier frequency an order of magnitude higher than those reported in familiar continental species, such as D. melanogaster (170 hertz). Drosophila fasciculisetae's song resembles a cicada's more than a fly's song. The song of D. cyrtoloma (Maui) has a complex pulse rhythm more typical of crickets than flies. The pulse song of D. silvestris (Hawaii) closely resembles that of D. melanogaster in both pulse rhythm and carrier frequency, but D. melanogaster sings by vibrating its wings, whereas D. silvestris sings through abdominal vibrations. These mechanisms are radical departures from the continental wing song mechanism and are further examples of the remarkable behavioral innovation that has occurred in the Drosophila of Hawaii during their evolutionary transit through these islands.

Animal Communication↗