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Masakado Kawata


kawata@m.tohoku.ac.jp

Journal articles

2010
J R Bridle, J Polechova, M Kawata, R K Butlin (2010)  Why is adaptation prevented at ecological margins? New insights from individual-based simulations   Ecology Letters 13: 485-494  
Abstract: All species are restricted in their distribution. Currently, ecological models can only explain such limits if patches vary in quality, leading to asymmetrical dispersal, or if genetic variation is too low at the margins for adaptation. However, population genetic models suggest that the increase in genetic variance resulting from dispersal should allow adaptation to almost any ecological gradient. Clearly therefore, these models miss something that prevents evolution in natural populations. We developed an individual- based simulation to explore stochastic effects in these models. At high carrying capacities, our simulations largely agree with deterministic predictions. However, when carrying capacity is low, the population fails to establish for a wide range of parameter values where adaptation was expected from previous models. Stochastic or transient effects appear critical around the boundaries in parameter space between simulation behaviours. Dispersal, gradient steepness, and population density emerge as key factors determining adaptation on an ecological gradient.
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M E Tsuda, M Kawata (2010)  Evolution of gene regulatory networks by fluctuating selection and intrinsic constraints.   PLoS Computational Biology 6: e1000873  
Abstract: Various characteristics of complex gene regulatory networks (GRNs) have been discovered during the last decade, e.g., redundancy, exponential indegree distributions, scale-free outdegree distributions, mutational robustness, and evolvability. Although progress has been made in this field, it is not well understood whether these characteristics are the direct products of selection or those of other evolutionary forces such as mutational biases and biophysical constraints. To elucidate the causal factors that promoted the evolution of complex GRNs, we examined the effect of fluctuating environmental selection and some intrinsic constraining factors on GRN evolution by using an individual-based model. We found that the evolution of complex GRNs is remarkably promoted by fixation of beneficial gene duplications under unpredictably fluctuating environmental conditions and that some internal factors inherent in organisms, such as mutational bias, gene expression costs, and constraints on expression dynamics, are also important for the evolution of GRNs. The results indicate that various biological properties observed in GRNs could evolve as a result of not only adaptation to unpredictable environmental changes but also non-adaptive processes owing to the properties of the organisms themselves. Our study emphasizes that evolutionary models considering such intrinsic constraining factors should be used as null models to analyze the effect of selection on GRN evolution.
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2009
Lázaro Miguel Echenique-Díaz, Jun Yokoyama, Osamu Takahashi, Masakado Kawata (2009)  Genetic structure of island populations of the endangered bat Hipposideros turpis turpis : implications for conservation   Population Ecology 51: 1. 153-160  
Abstract: Efforts for the conservation of the endangered bat species Hipposideros turpis turpis in southern Japan are hampered by a lack of information about its biology and natural history and by the increasing effect of human activities. In an attempt to address some of the conserva- tion challenges faced by this species, we studied the genetic structure and dispersal of intra- and interisland populations using six species-specific microsatellite mark- ers. In particular, we sought to establish the relationship between island populations and to define effective man- agement units for conservation. Pairwise co-ancestry index (FST) analysis, analysis of molecular variance, and Bayesian clustering suggested the presence of significant genetic differentiation between islands but little differen- tiation within them. The small Yonaguni Island population appeared to be not only geographically isolated, but also genetically isolated. This population is at the greatest risk of extinction, considering its size and low genetic variation. The larger populations on Iriomote and Ishigaki Islands are genetically related to each other to a greater degree and exhibit higher genetic variation than the Yonaguni Island population. This suggests that these two island populations should be included in a single management unit, while bats from Yonaguni Island should be managed independently and given higher priority for conservation. Actions such as defining vegetation corridors between colonies, as well as building gates at the entrance of the largest known colony, should be included in the conservation agenda of this still poorly known species.
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Akiko N Mizuno, Masakado Kawata (2009)  The effects of the evolution of stoichiometry-related traits on population dynamics in plankton communities   Journal of Theoretical Biology 259: 209-218  
Abstract: Both ecological stoichiometry and the evolution of traits for energetic interactions such as prey protection and predatory efficiency are considered to be important aspects affecting population dynamics. However, no attempt has been made to examine the effect of the evolution of traits relating to stoichiometry. This study first examined the effects of the evolution of nutrient utilization traits (i.e., the minimum nutrient content of prey, the maximum nutrient uptake affinity of prey and the nutrient contents of predators) on population dynamics in a plankton community. When the evolution of these traits was assumed, the range of the nutrient loading conditions where the system became unstable was smaller than when the evolution was not assumed, but the range of the conditions for zooplankton extinction became larger. Furthermore, when the trade-offs (i.e. genetic correlation between the traits) were assumed, the system rarely became extinct and the range of the nutrient loading conditions where the system became stable became larger through evolution. Stable dynamics were caused by increasing uptake affinity through evolution, and zooplankton extinction was caused by decreasing the minimum content of limiting nutrients. Thus, our results suggest that the evolution of traits relating to stoichiometry can affect the dynamics of the systems, and the outcomes the dynamics change greatly depend on which traits can evolve.
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Yosuke Kodaira, Hajime Ohtsuki, Jun Yokoyama, Masakado Kawata (2009)  Size-dependent foraging gene expression and behavioral caste differentiation in Bombus ignitus   BMC Research Notes 2: 184  
Abstract: Background: In eusocial hymenopteran insects, foraging genes, members of the cGMP-dependent protein kinase family, are considered to contribute to division of labor through behavioral caste differentiation. However, the relationship between foraging gene expression and behavioral caste in honeybees is opposite to that observed in ants and wasps. In the previously examined eusocial Hymenoptera, workers behave as foragers or nurses depending on age. We reasoned that examination of a different system of behavioral caste determination might provide new insights into the relationship between foraging genes and division of labor, and accordingly focused on bumblebees, which exhibit size-dependent behavioral caste differentiation. We characterized a foraging gene (Bifor) in bumblebees (Bombus ignitus) and examined the relationship between Bifor expression and size-dependent behavioral caste differentiation. Findings: A putative open reading frame of the Bifor gene was 2004 bp in length. It encoded 668 aa residues and showed high identity to orthologous genes in other hymenopterans (85.3-99.0%). As in ants and wasps, Bifor expression levels were higher in nurses than in foragers. Bifor expression was negatively correlated with individual body size even within the same behavioral castes (regression coefficient = -0.376, P < 0.001, all individuals; -0.379, P = 0.018, within foragers). Conclusion: These findings indicate that Bifor expression is size dependent and support the idea that Bifor expression levels are related to behavioral caste differentiation in B. ignitus. Thus, the relationship between foraging gene expression and behavioral caste differentiation found in ants and wasps was identified in a different system of labor determination.
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2008
Hajime Yoshino, Kyle N Armstrong, Masako Izawa, J Yokoyama, Masakado Kawata (2008)  Genetic and acoustic population structuring in the Okinawa Least Horsehoe bat: inter-colony acoustic differnces maintained by vertical maternal transmission?   Molecular Ecology 17: 23. 4978-4991  
Abstract: The origin and meaning of echolocation call frequency variation within rhinolophid bats is not well understood despite an increasing number of allopatric and sympatric examples being documented. A bimodal distribution of mean regional call frequency within the Okinawa-jima Island population of Rhinolophus cornutus pumilus (Rhinolophidae) provided a unique oppor- tunity to investigate geographic call frequency variation early in its development. Individual resting echolocation frequencies, partial mitochondrial DNA D-loop sequences and genotypes from six microsatellite loci were obtained from 288 individuals in 11 colonies across the entire length of the island, and nearby Kume-jima Island. Acoustic differences (5–8 kHz) observed between the north and south regions have been maintained despite evidence of sufficient nuclear gene flow across the middle of the island. Significant subdivision of maternally inherited D-loop haplotypes suggested a limitation of movement of females between regions, but not within the regions, and was evidence of female philopatry. These results support a ‘maternal transmission’ hypothesis whereby the difference in the constant frequency (CF) component between the regions is maintained by mother–offspring transmission of CF, the restricted dispersal of females between regions and small effective population size. We suggest that the mean 5–8 kHz call frequency difference between the regions might develop through random cultural drift.
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Hajime Ohtsuki, Jun Yokoyama, Nobuyoshi Ohba, Yoshihiro Ohmiya, Masakado Kawata (2008)  Nitric oxide synthase (NOS) in the Japanese fireflies Luciola lateralis and Luciola cruciata   Archives of Insect Biochemistry and Physiology 69: 189-198  
Abstract: Species-specific flash patterns in firefly species are important for the investigation of the evolution of Lampyridae. Since nitric oxide synthase (NOS) is one of the key enzymes controlling flash patterns, we determined the cDNA sequences of NOS in the Japanese fireflies Luciola lateralis and L. cruciata. The identity of the NOS sequences was very high between these 2 species. Firefly NOS also exhibited a high identity with those of other insect species, and the cofactor-binding domains were particularly well conserved. Many negatively selected sites were detected throughout the NOS sequences; however, no positive selection was detected. The phylogenetic relationship of insect NOS was different from that of the general classification system, although the lineages corresponded to the major recognized taxonomic groups.L
Notes: NO synthase; cDNA cloning; molecular evolution; Luciola lateralis; Luciola cruciata
2007
Masaru Imai, Yu-ichiro Tago, Makoto Ihara, Masakado Kawata, Kazuo Yamamoto (2007)  Role of the 5´ → 3´ exonuclease and Klenow fragment of Escherichia coli DNA polymerase I in base mismatch repair   Molecular Genetics and Genomics 278: 211-220  
Abstract: We have previously demonstrated that the Escherichia coli strain mutS ΔpolA had a higher rate of transition and minus frameshift mutations than mutS or ΔpolA strains. We argued that DNA polymerase I (PolI) corrects transition mismatches. PolI, encoded by the polA gene, possesses Klenow and 5´ → 3´ exonuclease domains. In the present study, rates of mutation were found to be higher in Klenow-defective mutS strains and 5´ → 3´ exonuclease-defective mutS strains than mutS or polA strains. The Klenow-defective or 5´ → 3´ exonuclease-defective mutS strains showed a marked increase in transition mutations. Sites of transition mutations in mutS, Klenow-defective mutS and 5´ → 3´ exonuclease-defective mutS strains are different. Thus, it is suggested that, in addition to mutS function, both the Klenow and 5´ → 3´ exonuclease domains are involved in the decrease of transition mutations. Transition hot and warm spots in mutS + polA + strains were found to differ from those in mutS and mutS ΔpolA strains. We thus argue that all the spontaneous transition mutations in the wild-type strain do not arise from transition mismatches left unrepaired by the MutS system or MutS PolI system.
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Ayako Shoji, Jun Yokoyama, Masakado Kawata (2007)  Molecular Phylogeny and Genetic Divergence of the Introduced Populations of Japanese Guppies, Poecilia reticulata   Conservation Genetics 8: 261-271  
Abstract: A phylogenetic tree of guppies from seven introduced Japanese populations was constructed using a mitochondrial gene, and the relationship between these Japanese populations and indigenous South American populations was examined. Phylogenetic trees constructed by maximum parsimony and neighbor joining methods indicated four major groups: Group A: Trinidadian populations; Group B: the northernmost Okinawa populations; Group C: the northern Okinawa populations; Group D: Shimoda populations and the remainder of Trinidadian, Venezuelan, and southern Okinawa populations. Considerable genetic differences were observed among the haplotypes within each Okinawa population similar to that found among haplotypes of different native populations, but not within the Shimoda population, because each Okinawa population included haplotypes belonging to different groups. Genetic differences between Groups B, C, and D might reflect those of the introduced lineages; however, these differences might not result from the divergence after introduction events. These results suggested that multiple introductions of different breeding lineages might cause considerable divergence within and between Okinawa populations
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Naoaki Yamamoto, Jun Yokoyama, Masakado Kawata (2007)  Relative resource abundance explains butterfly biodiversity in island communities   Proceedings of the National Academy of Sciences of the United States of America 104: 10524-10529  
Abstract: Ecologists have long been intrigued by the factors that control the pattern of biodiversity, i.e., the distribution and abundance of species. Previous studies have demonstrated that coexisting species partition their resources and/or that the compositional similarity between communities is determined by environmental factors, lending support to the niche-assembly model. However, no attempt has been made to test whether the relative amount of resources that reflects relative niche space controls relative species abundance in communities. Here, we demonstrate that the relative abundance of butterfly species in island communities is significantly related to the relative biomasses of their host plants but not to the geographic distance between communities. In the studied communities, the biomass of particular host plant species positively affected the abundance of the butterfly species that used them, and consequently, influenced the relative abundance of the butterfly communities. This indicated that the niche space of butterflies (i.e., the amount of resources) strongly influences butterfly biodiversity patterns. We present this field evidence of the niche-apportionment model that propose that the relative amount of niche space explains the pattern of the relative abundance of the species in communities.
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Satoshi Kato, Jotaro Urabe, Masakado Kawata (2007)  Effects of temporal and spatial heterogeneities created by consumer-driven nutrient recycling on algal diversity   Journal of Theoretical Biology 245: 364-377  
Abstract: A spatially explicit plant–herbivore model composed of planktonic herbivores, algal preys and nutrients was constructed to examine the effects of consumer-driven nutrient recycling (CNR) on the algal species richness with and without spatial structure. The model assumed that either of two essential nutrients (N and P) limited growth of algal populations and that consumer individuals moved randomly in the lattice and grazed all the algal species with the same efficiency. The results showed that when there was no CNR, the number of persistent algal species was affected by neither supply rates of external nutrients nor spatial structure and was consistently low. When consumers recycled nutrients according to their stoichiometry, the algal species richness changed with supply rates of external nutrients depending on spatial structure: the algal species richness decreased with increasing nutrient loadings when there were no spatial structure because CNR increased the probability of stochastic extinction of algal species by amplifying the oscillation of algae-consumer dynamics. However, when spatial structures were created by the migration of consumers, CNR increased the algal species richness in a range of nutrient loadings because spatial variation of grazing pressure functioned to stabilize the algal-consumer dynamics. The present study suggests that through grazing and nutrient recycling, consumer individuals can create ephemeral heterogeneity in growth environments for algal species and that this ephemerality is one of the keys to understanding algal species in nature.
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Masakado Kawata, Ayako Shoji, Shoji Kawamura, Ole Seehausen (2007)  A genetically explicit model of speciation by sensory drive within a continuous population in aquatic environments   BMC Evolutionary Biology 7: 99  
Abstract: Background The sensory drive hypothesis predicts that divergent sensory adaptation in different habitats may lead to premating isolation upon secondary contact of populations. Speciation by sensory drive has traditionally been treated as a special case of speciation as a byproduct of adaptation to divergent environments in geographically isolated populations. However, if habitats are heterogeneous, local adaptation in the sensory systems may cause the emergence of reproductively isolated species from a single unstructured population. In polychromatic fishes, visual sensitivity might become adapted to local ambient light regimes and the sensitivity might influence female preferences for male nuptial color. In this paper, we investigate the possibility of speciation by sensory drive as a byproduct of divergent visual adaptation within a single initially unstructured population. We use models based on explicit genetic mechanisms for color vision and nuptial coloration. Results We show that in simulations in which the adaptive evolution of visual pigments and color perception are explicitly modeled, sensory drive can promote speciation along a short selection gradient within a continuous habitat and population. We assumed that color perception evolves to adapt to the modal light environment that individuals experience and that females prefer to mate with males whose nuptial color they are most sensitive to. In our simulations color perception depends on the absorption spectra of an individual's visual pigments. Speciation occurred most frequently when the steepness of the environmental light gradient was intermediate and dispersal distance of offspring was relatively small. In addition, our results predict that mutations that cause large shifts in the wavelength of peak absorption promote speciation, whereas we did not observe speciation when peak absorption evolved by stepwise mutations with small effect. Conclusion The results suggest that speciation can occur where environmental gradients create divergent selection on sensory modalities that are used in mate choice. Evidence for such gradients exists from several animal groups, and from freshwater and marine fishes in particular. The probability of speciation in a continuous population under such conditions may then critically depend on the genetic architecture of perceptual adaptation and female mate choice.
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2006
Toru Usami, Jun Yokoyama, Kohei Kubota, Masakado Kawata (2006)  Genital lock-and-key system and premating isolation by mate preference in carabid beetles (Carabus subgenus Ohomopterus)   Biological Journal of the Linnean Society 87: 145-154  
Abstract: The shapes and lengths of copulatory pieces and vaginal appendices of the carabid beetle subgenus Ohomopterus (genus Carabus) vary among species. In Japan, the species in the group with a medium body size (C. yaconinus, C. iwawakianus, C. maiyasanus, C. uenoi, C. arrowianus, C. esakii, and C. insulicola) are usually allopatric or parapatric, except at Mt Kongosan, where C. uenoi, C. iwawakianus, and C. yaconinus are sympatrically distributed. The degree of premating isolation by mate preference was high between sympatric populations, irrespective of the genetic distance between them. However, premating isolation was absent between parapatric populations. The degree of premating isolation for allopatric populations spanned a wide range of isolation values. Thus, mate discrimination by males seems to have evolved mostly between sympatric pairs. These results suggest two hypotheses. First, premating isolation has evolved through reinforcement or through reproductive character displacement after sympatric contact. Second, premating isolation has evolved in allopatry, and as a result of premating isolation, the species can coexist in sympatry. We also examined the degree of mechanical isolation between C. uenoi and C. iwawakianus (a sympatric pair), which have a very large difference in the length of the copulatory piece. The insertion success was low and only one female produced viable offspring among 15 crosses; however, death in females due to copulation was rare. For sympatric matings between C. uenoi and C. iwawakianus, a large difference in the genital size might reduce the gene flow with small mating costs. Gene flow that was significantly reduced by genital difference might cause either the evolution of premating isolation through reinforcement/reproductive character displacement or through the maintenance of a high degree of premating isolation following sympatric contact.
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Takehiko I Hayashi, Masakado Kawata (2006)  Impact of ancestral populations on postzygotic isolation in allopatric speciation   Population Ecology 48: 121-130  
Abstract: Postzygotic isolation evolves due to an accumulation of substitutions (potentially deleterious alleles in hybrids) in populations that have become geographically isolated. These potentially deleterious alleles might also be maintained in ancestral populations before geographic isolation. We used an individual-based model to examine the effect of the genetic state of an ancestral population on the evolution of postzygotic isolation after geographic isolation of a population. The results showed that the number of loci at which degenerative alleles are fixed in an ancestral population at equilibrium significantly affects the evolutionary rates of postzygotic isolation between descendant allopatric populations. Our results suggest that: (1) a severe decrease in population size (e.g., less than ten individuals) is not necessary for the rapid evolution of postzygotic isolation (e.g., <10,000 generation); (2) rapid speciation can occur when there is a large difference in the equilibrium number of accumulated degenerative alleles between ancestral and descendant populations; and (3) in an ancestral population maintained at a small effective population size for a long period of time, postzygotic isolation rarely evolves if back mutations that restore the function of degenerative alleles are limited.
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2005
Yuki Nagata, Genta Kawaguchi, Yu-ichiro Tago, Masaru Imai, Takashi Watanabe, Shigehisa Sakurai, Makoto Ihara, Masakado Kawata, Kazuo Yamamoto (2005)  Absence of strand bias for deletion mutagenesis during chromosomal leading and lagging strand replication in Escherichia coli   Genes & Genetic Systems 80: 1-8  
Abstract: Investigations were carried out to determine whether both DNA strands involved in Escherichia coli chromosomal DNA replication are replicated with similar accuracy. Experiments consisted of measuring the forward mutation rate from tonB+ to tonB– in pairs of polA deficient strains in which the chromosomal target gene tonB was oriented in the two possible directions relative to the origin of replication, oriC. Within these pairs, the tonB sequence would be subjected to leading strand replication in one orientation and to lagging strand replication in the other. The most common tonB mutations in the polA1 strain were deletions followed by frameshifts. Among the deletions, a strong hotspot site with a 13-base deletion in the polA1 strains accounted for 18 of the 33 deletions in the one orientation, and 31 of the 58 deletions in the other. The results suggested that the two strands were replicated with equal or similar accuracy for deletion formation.
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Noe Matsushima, Masakado Kawata (2005)  The choice of oviposition site and the effects of density and oviposition timing on survivorship in Rana japonica   Ecological Research 20: 81-87  
Abstract: In species that lack parental care and whose larvae are restricted to a given space throughout development, choice of egg-laying or reproductive sites often greatly influences fitness. Furthermore, reproductive success of individuals may be affected not only by conditions of the breeding area but also by the time when they start to breed, because intraspecific variation in the timing of breeding may result in asymmetric competition between larvae. We addressed these issues in an experimental and field study of the Japanese brown frog, Rana japonica. We tested whether the survival success of larvae decreased with an increase in the number of deposited egg masses and with a greater delay in oviposition. We found that the number of egg masses and timing of oviposition together significantly predicted larval survival per clutch. Moreover, we observed the natural oviposition of R. japonica to examine the prediction that if the density of larvae and the timing of oviposition affect survivorship of larvae, R. japonica will avoid depositing eggs in pools in which egg masses have already been deposited. We found that for small pools only, R. japonica tended to favor oviposition at unoccupied pools (those lacking egg masses and larvae) in the spawning season. For large pools, however, adults favored those already occupied by egg masses in the spawning period. The density of egg masses and tadpoles or the timing of oviposition may have less effect on spawning or the survival of tadpoles in large pools than in small pools.
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Masakado Kawata, Hiroshi Sawada, Jun Yokoyama, Lázaro M Echenique-Díaz, Yasuyuki Ishibashi (2005)  The effects of spatial habitat structure on the evolution of density-dependent growth and reproduction in freshwater snails   Hydrobiologia 533: 229-241  
Abstract: We examined the growth and reproductive rates of freshwater snails, Physa acuta, in two habitat types. In the Asabata habitat, snails lived in isolated water pools, which occasionally joined to form a single large pool; in the Kakegawa habitat, they lived in a slow-running water way. Genetic structure assessments using three microsatellite loci supports the idea that a stable panmictic population occupies the Kakegawa habitat. The Asabata habitat, however, is occupied with an alternate mixing population as revealed by microsatellite data. The Asabata population might alternate between localized mating within isolated pools (as revealed by high F IS and F IT values) when the water levels are low and panmixia (as revealed by the low F ST values and AMOVA analysis) when the habitat is flooded. Laboratory experiments, using snails collected from the two habitats, showed that juvenile snails grew faster, laid more eggs, and laid them earlier in the Asabata habitat than in the Kakegawa habitat. Growth rates were lower at high density than at low density in the Kakegawa habitat; the inverse was true in the Asabata habitat. Density-dependent response of individual snail reproduction was higher in the Kakagawa habitat than in the Asabata habitats. The results support the hypothesis that spatial structure affects the evolution of density-dependent growth rates and of timing for reproduction.
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Masayoshi Tamura, Jun Yokoyama, Nobuyoshi Ohba, Masakado Kawata (2005)  Geographic differences in flash intervals and pre-mating isolation between populations of the Genji firefly, Luciola cruciata   Ecological Entomology 30: 241-245  
Abstract: 1. Two types of Genji firefly, Luciola cruciata, have been recognised on the basis of the inter-flash interval. 2. The fast-flash and slow-flash type fireflies, distributed in the western and eastern parts of Japan, exhibit a 2- and 4-s interval of bioluminescence flash, respectively. In addition, an intermediate-flash type is found on the boundary between these two regions. 3. The differences in the responses of the males of four field populations to different intervals of artificial flashes were examined. 4. The results revealed that in the Aomori and Sendai populations (slow-flash type), a significantly larger number of males approached the 4- and 5-s flash intervals than approached the 2- and 3-s intervals of artificial flashes. On the other hand, in the Ohtsu population (fast-flash type), a significantly larger number of males approached the 2- and 3-s flash intervals than the 4- and 5-s intervals of artificial flashes. The Inuyama population (intermediate-flash type) did not show any particular preferences to flash intervals. 5. This study shows that in firefly populations, geographic variation in flash pattern can cause some degree of pre-mating isolation between populations.
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2004
Osamu K Mikami, Masakado Kawata (2004)  Does interspecific territoriality reflect resource competition?: A theoretical model for the interspecific territoriality   Evolutionary Ecology Research 6: 765-775  
Abstract: In many studies, interspecific territorial behaviours among co-existing species have been used to infer the presence and the intensity of underlying ecological interactions between species, mainly resource competition. However, the theoretical background of this inference is insufficient. Hence, we constructed a simple theoretical model of interspecific territoriality assuming that interspecific territorial defence is the optimal behaviour. We discuss the factors promoting interspecific territoriality and the relationship between interspecific territoriality and ecological interactions. The model predicts that: (1) a territory holder preferentially excludes intruders of species with high ‘exclusion efficiency’; (2) the decision by the territory holder to exclude a certain species or not does not depend on the probability of finding intruders of the species or on the number of intrusions by the species; and (3) interspecific territoriality does not always reflect the intensity of ecological interactions between species. These results indicate that the observation of interspecific territoriality does not necessarily indicate the intensity of ecological interactions. In addition, if territory holders defend their territories as predicted by the present optimal model, the co-existence of competing species is promoted.
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Osamu K Mikami, Masanori Kohda, Masakado Kawata (2004)  A new hypothesis for species coexistence: male–male repulsion promotes coexistence of competing species   Population Ecology 46: 213-217  
Abstract: We propose a new hypothesis for species coexistence by considering behavioral interactions between individuals. The hypothesis states that repulsive behavior between conspecific males (male–male repulsion) creates space for competing species, which promotes their coexistence. This hypothesis can explain the coexistence of two competing species even when their ecological niches completely overlap in spatially homogeneous environments. In addition, the mechanisms underlying such behavior might play a role in enabling the coexistence of two species immediately after speciation, with little or no niche differentiation, as in the case of cichlid fish communities, for example. Although there is limited evidence supporting this hypothesis, it can nevertheless explain the occurrence of species coexistence and biodiversity, which cannot be explained by previous theories.
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2003
Y Nagata, M Kawata, J Komura, T Ono, K Yamamoto (2003)  X-ray-induced mutaions in Escherichia coli K-12 strains with altered DNA polymerase I activities.   Mutation Research/Fundamental and Molecular Mechanisms of Mutagenesis 528: 93-103  
Abstract: Spectra of ionizing radiation mutagenesis were determined by sequencing X-ray-induced endogenous tonB gene mutations in Escherichia coli polA strains. We used two polA alleles, the polA1 mutation, defective for Klenow domain, and the polA107 mutation, defective for flap domain. We demonstrated that irradiation of 75 and 50 Gy X-rays could induce 3.8- and 2.6-fold more of tonB mutation in polA1 and polA107 strains, respectively, than spontaneous level. The radiation induced spectrum of 51 tonB mutations in polA1 and 51 in polA107 indicated that minus frameshift, A:T→T:A transversion and G:C→T:A transversion were the types of mutations increased. Previously, we have reported essentially the same X-ray-induced tonB mutation spectra in the wild-type strain. These results indicate that (1) X-rays can induce minus frameshift, A:T→T:A transversion and G:C→T:A transversion in E. coli and (2) presence or absence of polymerase I (PolI) of E. coli does not have any effects on the process of X-ray mutagenesis.
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H Teshima, D Angus, Y Kuwahara, J Yokoyama, S Chiba, T Fukuda, H Ogimura, M Kawata (2003)  The evolution of extreme shell shape variation in the land snail Ainohelix editha: a phylogeny and hybrid zone analysis.   Molecular Ecology 12: 1869-1878  
Abstract: Ainohelix editha from Hokkaido, Japan, exhibit great geographical variation in their shell morphology. In particular, A. editha in two quite separate locations, Shimamaki and Samani, are striking because they are extremely flat and have a sharp keel, whereas at adjacent sites the shells are globular or depressed-globular. We used mitochondrial 16S rRNA and nuclear ITS-2 sequences to infer a phylogeny among 47 snails from 29 locations. Snails from the two keeled-flat populations clustered separately in the phylogeny, suggesting that this unusual shell form could have evolved independently. A morphological analysis of shells collected along a transect between keeled-flat and globular snail sites showed a cline for shell shape and the angle of the keel. Two different mtDNA lineages were found across the transect, with a cline for an ITS-2 single nucleotide polymorphism. Together, the results may suggest a lack of reproductive isolation between keeled-flat and globular snails, with possible introgression by hybridization
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2002
Masakado Kawata (2002)  Invasion of vacant niches and subsequent sympatric speciation   Proceedings of the Royal Society of London B 269: 55 - 63  
Abstract: An individual-based simulation study was conducted to examine the population dynamics of ‘invasion of a vacant niche’ and subsequent speciation (by reproductive isolation) when food resources are randomly distributed spatially within the habitat and the frequencies of different food types are bimodally distributed (i.e. smaller and larger sizes of food being most abundant). The initially vacant niche was that of unused larger sizes of food. When phenotypic variation for resource use (i.e. food sizes) was small in the initial population, and each female could choose a mate from anywhere in the habitat, the population could not invade the vacant niche. But when the dispersal distance of the offspring and the area within which a female could choose a mate were small (i.e. the genetic neighbourhood size was small), the population could, in most cases, evolve to use both smaller and larger food sizes and form sister species sympatrically, with each species utilizing one of the two niches (small and large sizes of food). When phenotypic variation in resource use in the initial population was large, the population could, in most cases, invade the vacant niche by evolving to use both smaller and larger sizes of food. The probability of speciation increased as the dispersal distance of offspring decreased. The results indicate that populations whose individuals have small Wright's genetic neighbourhoods may often exploit a vacant niche and diversify sympatrically in the process.
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2001
Masakado Kawata (2001)  The influence of neighborhood size and habitat shape on the accumulation of deleterious mutations   Journal of Theoretical Biology 211: 187-199  
Abstract: To examine the impact of genetic neighborhood size and habitat shape on genetic load and the accumulation of deleterious mutation, individual-based simulations were performed in continuously distributed habitats. The risk of extinction increased as both the area of the habitat and the neighborhood size decreased. When the neighborhood area became smaller than the habitat area, habitat shape also began to influence the risk of extinction by mutation loads, expected time to extinction being shorter in longer and narrower habitats than in a square habitat. Both the number of homozygous deleterious loci per individual and the mutation load in the population increased as the neighborhood size and total population size decreased. Neighborhood size and total population size both independently affected the average number of homozygous deleterious loci per individual. In addition, as the ratio of the long to the short side of the rectangle of a habitat increased, the average number of homozygous deleterious loci increased. When the areas of the habitats were held constant, the average number of homozygous loci and the mutation loads were smallest for a regular square and largest for the longest, narrowest habitat. These results suggest that the spatial genetic structure of an individual is an important factor in the accumulation of deleterious mutations and the risk of extinction by mutation meltdown.
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2000
Masakado Kawata, Jin Yoshimura (2000)  Speciation by sexual selection in hybridizing poplations without viability selection   Evolutionary Ecology Research 2: 897-909  
Abstract: We propose a new speciation mechanism for hybridizing populations based on frequency- dependent sexual selection theory. In contrast to previous reinforcement and symmetric speciation models, this model does not require viability selection for male sexual traits, selection against unfit hybrids or disruptive selection for ecological traits. The model assumes that females exhibit mating preferences based on male secondary sexual characters. Both female mating preferences and male secondary sexual traits are independent quantitative genetic traits. Females choose using one of five tactics. All individuals, including the hybrids, have the same ability to survive and reproduce. Two populations that differ only in their average values of female preference and male sexual traits are assumed to come into contact and hybridize. Most cases of the model frequently result in complete prezygotic isolation. The likelihood of speciation decreases with the degree of phenotypic overlap between the two merging popula- tions. However, the probability of speciation is not zero even when the two merging populations are identical
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Takayuki Gotoh, Masakado Kawata (2000)  The effects of spatial habitat structure on population variability of freshwater snails   Hydrobiologia 429: 157–169  
Abstract: Experiments were conducted to examine the effects of a habitat's spatial structure on population variability in two species of freshwater snails (Physa acuta and Austropeplea ollula). To alter the spatial structure of the habitat, vinyl chloride plates were hung in experimental tanks, providing three types of spatial structure: Complex structure, Simple structure and Control (no structure). In Experiment 1, the average number of individuals in a tank did not differ among the three types of structure 2 months after the introduction of the snails, but the variability of the number of individuals in the Complex structure tanks was lowest, whereas the variability in the Control tanks was highest. In Experiment 2, in addition to the spatial structure of the habitat, three types of species interaction were designed as experimental treatments: only P. acuta was introduced into the tanks (P. acuta tanks), only A. ollula was introduced into the tanks (A. ollulatanks) and both P. acuta and A. ollula were introduced into the tanks (two-species tanks). For the P. acuta tanks, the variability of the number of P. acuta individuals in the Complex structure tanks was lowest, and the variability in the Control tanks was highest when the effect of the number of individuals in a tank was subtracted. For the A. ollula tanks and the two-species tanks, there were no significant differences in the variability of the population size among the different treatments of spatial structure. The spatial distribution of P. acuta was more uniform than the distribution of A. ollula on the plates of complex structure. Our results indicate that the spatial structure of the habitat influences the variability of population size (the variance of the number of individuals in different populations during the earlier period after the introduction of the snails), but the effects depend on the spatial behavior of individuals and the interaction with other species
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1999
Masakado Kawata, Hiroko Agawa (1999)  Perceptual Scales of spatial heterogeneity of periphyton for freshwater snails   Ecology Letters 2: 210-214  
Abstract: The hypothesis that individuals recognize their environment as homogeneous when the scales of spatial heterogeneity of resources are smaller than a certain scale, but can distinguish them as patches when they are larger than that scale was tested using freshwater snails (Physa acuta) in various distributions in periphyton environments. In a pattern of periphyton distribution in which the size of algal cells was 47 mm, individuals moved significantly more slowly on algal cells than on nonalgal cells. However, in other patterns in which the sizes of algal cells were 23.5 mm and 15.7 mm, the speeds of individuals on algal and nonalgal cells were not significantly different. These results support the hypothesis that individuals use the environment homogeneously when the scales of spatial heterogeneity of resources are smaller than a certain scale, but they can distinguish between patches when the scales are larger.
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Masakado Kawata (1999)  The effect of dispersal behavior in group selection   Evolutionary Ecology Research 1: 663-680  
Abstract: Individual-based simulations were conducted to examine the spread of an altruistic allele by group selection. For the simulations, individuals with one locus determining altruistic behaviour were created in an environment with 13 patches. Individual fitness was determined by the number of altruists and the number of total individuals in a patch. Offspring disperse within the natal patch or into neighbouring patches depending on random values with a normal distribution having a zero mean and standard deviation σd (random dispersal model). The probability of the spread of the altruistic allele increased with increasing dispersal distance until a moderately low dispersal distance (σd = 50); thereafter, the probability decreased with increasing dispersal distance. A density-dependent dispersal model in which female offspring tend to disperse to the patch with lowest densities among five candidate dispersal sites was also tested. For the density-dependent dispersal model, the chance of spreading the altruistic allele was higher than that of the random dispersal model. The results can be explained by two components of group selection: the extinction of groups with lower frequencies of altruists (selective extinction) and higher frequencies of migrants of altruists due to high productivities of altruistic groups (effective migrants). Evolution of altruism by interdemic group selection only (i.e. by differential extinctions and recolonization of demes) appears unlikely, since it requires high rates of both extinction and recolonization with low migration among surviving demes. However, the present study showed that moderately low migration can create favourable conditions under which both selective extinction and effective migrants simultaneously increase the probability of spreading the altruistic alleles
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1998
1997
Masakado Kawata (1997)  Exploitative competition and ecologically effective abundance   Ecological Modelling 97: 125-137  
Abstract: This study analyzes the effect of exploitative competition and density dependence by using an individual-based simulation model. There are two components of exploitative competition (leftovers and resource-recovery competition). In leftovers competition, individuals compete for food resources that have been left by others. In resource-recovery competition, the consumption of foods by individual predators affects growth rates of food populations. An individual-based simulation was conducted to examine how these two types of competition affect density-dependent processes. In the simulation, individual predators search for food organisms within their home range. Food organisms reproduce in three different ways: replacement, recruitment and neighborhood recruitment. In replacement, all food organisms die and are replaced with a new food population. In recruitment, food organisms do not disappear during simulation unless they are eaten by predators, and new food organisms are recruited based on their density. In neighborhood recruitment, food organisms reproduce depending on their abundance within the neighborhood area. The simulation was conducted by varying the number of predators, the home range size of individual predators, the area of simulation, the reproductive rates of food organisms, and the reproductive modes of the food organisms. The number of food organisms taken by an individual predator, and the number of predators over several different scales, were measured. A multiple regression was conducted to examine the relationships between several different estimations of predator abundance and the number of food organisms taken by an predator. The effects of different types of exploitative competition differed depending upon the spatial scale or estimated population density. The number of predators within the home range of an individual predator is negatively related to individual food gain. This suggests that leftovers competition can be detected within the home range of an individual predator. In the case in which the food population was regulated over the whole simulation area, the number of predators in the whole area was positively related to food gain of individual predator in the case of recruitment and neighborhood recruitment of reproduction of food, but positively in replacement. In the case in which a food population was regulated in smaller parts of the whole area, the estimated number of predators within a food population were related to food gain of an individual predator but the total population size does not affect the food gain.
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Hiroki Hinoue, Masakado Kawata (1997)  The effect of neighborhood densities on reproductive success in a heronry   Ecoscience 4: 465-469  
Abstract: The effects of the number of nests within a neighboring area on reproductive success were examined in Egretta garzetta and Ardea cinerea in a heronry. Two neighboring areas were defined: (1) the Neighborhood is an area enclosed by a circle of radius N whose center is located at a focal nest; (2) the Surrounding is an area enclosed by a circle of radius S (S>N) that excludes the Neighborhood. N and S were set at several different values. The number of nests in the Neighborhood had a significant negative effect on reproductive success when N=0.5 m and S=1 m in E. garzetta and N=1.5 m and S=4.5 m. The number of nests within the Surrounding had a significant positive effect on the reproductive success of A. cinerea when N=1.5 m and N=3 m or S=4.5 m. These results suggest that to examine individual differences in reproductive success within colonies, we should choose the appropriate Neighborhood and Surrounding to detect the effects of interactions among neighboring individuals
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Masakado Kawata (1997)  The loss of genetic variabilities in a fragmented continuous population   Researches on Population Ecology 39: 227-237  
Abstract: An individual-based simulation model was used to examine the effect of population subdivision, dispersal distance of offspring, and migration rates between subpopulations on genetic variability (HI , HS and HT) in a continuously distributed population. Some difficulties with mathematical models of a continuously distributed population have been pointed out. The individual-based model can avoid these difficulties and can be used to examine genetic variability in a population within which individuals are distributed continuously and in which the dispersal of individuals is disturbed by geographical or artificial barriers. The present simulation showed that the pattern of decrease in HI had three stages. During the first stage, HI decreased with the rates predicted by Wright's neighborhood size. During the second stage, HI decreased more rapidly when the migration rate decreased, while during the third stage, it decreased less rapidly when the migration rate decreased. Increasing the number of subdivisions increased the rate of decrease after the 200th generation. The pattern of decrease in HT was classified into 2 stages. During the first stage, the rates of decrease corresponded with those of a randomly mating population. During the second stage, a decrease in the migration rates of the subpopulations slowed the rate of decrease in HT. A uniform spatial distribution and a reduced total dispersal distance of offspring caused HI, HS, and HT to decrease more rapidly. Habitat fragmentation in a continuously distributed population usually was detrimental to the genetic variability in the early generations. Other implications of the results for conservation are discussed.
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Book chapters

2003

Book Review

2004
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