Researcher Database

Masayo Soma
Faculty of Science Biological Sciences Behavioral Neuroethology
Associate Professor

Researcher Profile and Settings

Affiliation

  • Faculty of Science Biological Sciences Behavioral Neuroethology

Job Title

  • Associate Professor

J-Global ID

Research Interests

  • 鳴禽類   性淘汰   コミュニケーション   求愛   進化   社会性   ディスプレイ   発達   歌鳥   比較認知   

Research Areas

  • Humanities & social sciences / Cognitive sciences
  • Life sciences / Basic brain sciences
  • Life sciences / Evolutionary biology

Academic & Professional Experience

  • 2010 - Today 北海道大学 理学(系)研究科(研究院) 理学(系)研究科(研究院) 准教授

Research Activities

Published Papers

  • Soma M, Iwama M, Nakajima R, Endo R
    Royal Society Open Science 6 190563  2019/06 [Refereed][Not invited]
  • Vocalization can mediate male-male sexual interactions in Java sparrows
    Hiroko Adachi, Masayo Soma
    Animal Biology 2018/12 [Refereed][Invited]
  • Ota N, Gahr M, Soma M
    Science Advances 4 (10) eaat4779  2018/10 [Refereed][Not invited]
  • Soma M, Garamszegi LZ
    Behavioral Ecology Oxford University Press ({OUP}) 2018/05 [Refereed][Not invited]
  • A. C. R. Gomes, C. Funghi, M. Soma, M. D. Sorenson, G. C. Cardoso
    JOURNAL OF EVOLUTIONARY BIOLOGY 30 (7) 1336 - 1346 1010-061X 2017/07 [Refereed][Not invited]
     
    Sexual traits (e.g. visual ornaments, acoustic signals, courtship behaviour) are often displayed together as multimodal signals. Some hypotheses predict joint evolution of different sexual signals (e.g. to increase the efficiency of communication) or that different signals trade off with each other (e.g. due to limited resources). Alternatively, multiple signals may evolve independently for different functions, or to communicate different information (multiple message hypothesis). We evaluated these hypotheses with a comparative study in the family Estrildidae, one of the largest songbird radiations, and one that includes many model species for research in sexual selection and communication. We found little evidence for either joint evolution or trade-offs between song and colour ornamentation. Some negative correlations between dance repertoire and song traits may suggest a functional compromise, but generally courtship dance also evolved independently from other signals. Instead of correlated evolution, we found that song, dance and colour are each related to different socio-ecological traits. Song complexity evolved together with ecological generalism, song performance with investment in reproduction, dance with commonness and habitat type, whereas colour ornamentation was shown previously to correlate mostly with gregariousness. We conclude that multimodal signals evolve in response to various socio-ecological traits, suggesting the accumulation of distinct signalling functions.
  • Masayo Soma, Midori Iwama
    PLOS ONE 12 (3) e0172655  1932-6203 2017/03 [Refereed][Not invited]
     
    Mutual interactions between sexes have multiple signalling functions. Duet singing in songbirds is related to mutual mate guarding, joint resource defence, and signalling commitment. Coordinated visual displays of mating pairs are thought to perform similar functions, but are less well understood. The current study evaluated mutual interactions in an Estrildid species to explore the relative importance of duet dancing and male singing in mating success of pairs in a first encounter. When Java sparrows (Lonchura oryzivora) court prospective mates, only males sing. However, both males and females perform courtship dances, often in a duet-like manner. These dances are typically terminated by female copulation solicitation displays (CSDs). In the current study, we observed higher mating success when courtship dances were mutually exchanged, and when males sang. However, the sex initiating the courtship did not affect mating success. Most females produced CSDs after duet dancing but before hearing the entire song, indicating that duet dancing played a crucial role in mating. This finding highlights an unexplored aspect of duetting behaviour in the process of mutual mate choice. These results conflict with the majority of past songbird research, which has interpreted songs as primary behavioural sexual signals.
  • Nao Ota, Manfred Gahr, Masayo Soma
    BIOACOUSTICS-THE INTERNATIONAL JOURNAL OF ANIMAL SOUND AND ITS RECORDING 26 (2) 161 - 168 0952-4622 2017 [Refereed][Not invited]
     
    Vocalizations have been elucidated in previous songbird studies, whereas less attention has been paid to non-vocal sounds.In the bluecapped cordon-bleu (Uraeginthus cyanocephalus), both sexes perform courtship displays that are accompanied by singing and distinct body movements (i.e.dance).Our previous study revealed that their courtship bobbing includes multiple rapid steps.This behaviour is quite similar to human tap dancing, because it can function as both visual and acoustic signals.To examine the acoustic signal value of such steps, we tested if their high-speed step movements produce non-vocal sounds that have amplitudes similar to vocal sounds. We found that step behaviour affected step sound amplitude. Additionally, the dancing step sounds were substantially louder than feet movement sounds in a non-courtship context, and the amplitude range overlapped with that of song notes.These results support the idea that in addition to song cordon-bleus produce acoustic signals with their feet.
  • Kentaro Yamada, Masayo Soma
    JOURNAL OF AVIAN BIOLOGY 47 (6) 865 - 870 0908-8857 2016/12 [Refereed][Not invited]
     
    Song is a notable sexual signal of birds, and serves as an honest indicator of male quality. Condition dependence of birdsong has been well examined from the viewpoint of the developmental stress hypothesis, which posits that complex songs assure fitness because learned acoustic features of songs are especially susceptible to early-life stress that young birds experience in song learning periods. The effect of early stress on song phenotypes should be crucial, especially in age-limited song learners which sing stereotyped songs throughout life. However, little attention has been paid to non-learned song features that can change plastically even in adulthood of age-limited song-learners. Although it has been shown that food availability affects song rate in wild songbirds, there is limited evidence of the link between favorable nutritional conditions and song phenotypes other than song rate. Under the prediction that singing behavior reflects an individual's recent life history, we kept adult Bengalese finch males under high-nutrition or normal diet for a short term, and examined changes in body mass and songs. We found that birds on a high-nutrition diet showed higher song output (e.g. song rate and length) compared with those of the control group, while changes in body mass were moderate. In addition, note repertoire became more consistent and temporal structures got faster in both nutrition and control groups, which indicates that songs were subject to other factors than nutrition. Considering that female estrildid finches, including Bengalese and zebra finches, show a preference toward complex songs as well as longer songs and higher song rate, it is plausible that different aspects of singing behavior signal different male qualities, and provide multifaceted clues to females that choose mates.
  • Nao Ota, Manfred Gahr, Masayo Soma
    SCIENTIFIC REPORTS 5 2045-2322 2015/11 [Refereed][Not invited]
     
    According to classical sexual selection theory, complex multimodal courtship displays have evolved in males through female choice. While it is well-known that socially monogamous songbird males sing to attract females, we report here the first example of a multimodal dance display that is not a uniquely male trait in these birds. In the blue-capped cordon-bleu (Uraeginthus cyanocephalus), a socially monogamous songbird, both sexes perform courtship displays that are characterised by singing and simultaneous visual displays. By recording these displays with a high-speed video camera, we discovered that in addition to bobbing, their visual courtship display includes quite rapid step-dancing, which is assumed to produce vibrations and/or presumably non-vocal sounds. Dance performances did not differ between sexes but varied among individuals. Both male and female cordon-bleus intensified their dance performances when their mate was on the same perch. The multimodal (acoustic, visual, tactile) and multicomponent (vocal and non-vocal sounds) courtship display observed was a combination of several motor behaviours (singing, bobbing, stepping). The fact that both sexes of this socially monogamous songbird perform such a complex courtship display is a novel finding and suggests that the evolution of multimodal courtship display as an intersexual communication should be considered.
  • Masayo Soma, Chihiro Mori
    PLOS ONE 10 (5) 1932-6203 2015/05 [Refereed][Not invited]
     
    Music and dance are two remarkable human characteristics that are closely related. Communication through integrated vocal and motional signals is also common in the courtship displays of birds. The contribution of songbird studies to our understanding of vocal learning has already shed some light on the cognitive underpinnings of musical ability. Moreover, recent pioneering research has begun to show how animals can synchronize their behaviors with external stimuli, like metronome beats. However, few studies have applied such perspectives to unraveling how animals can integrate multimodal communicative signals that have natural functions. Additionally, studies have rarely asked how well these behaviors are learned. With this in mind, here we cast a spotlight on an unusual animal behavior: nonvocal sound production associated with singing in the Java sparrow (Lonchura oryzivora), a songbird. We show that male Java sparrows coordinate their bill-click sounds with the syntax of their song-note sequences, similar to percussionists. Analysis showed that they produced clicks frequently toward the beginning of songs and before/after specific song notes. We also show that bill-clicking patterns are similar between social fathers and their sons, suggesting that these behaviors might be learned from models or linked to learning-based vocalizations. Individuals untutored by conspecifics also exhibited stereotypical bill-clicking patterns in relation to song-note sequence, indicating that while the production of bill clicking itself is intrinsic, its syncopation appears to develop with songs. This paints an intriguing picture in which non-vocal sounds are integrated with vocal courtship signals in a songbird, a model that we expect will contribute to the further understanding of multimodal communication.
  • Soma M, Garamszegi LZ
    Frontiers in Ecology and Evolution 3 4  2015/02 [Refereed][Not invited]
  • Nao Ota, Masayo Soma
    JOURNAL OF AVIAN BIOLOGY 45 (6) 566 - 573 0908-8857 2014/11 [Refereed][Not invited]
     
    Birdsong is a sexual signal that serves as an indicator of male quality. There is already abundant evidence that song elaboration reflects early life-history because early developmental stress affects neural development of song control systems, and leaves irreversible adverse effects on song phenotypes. Especially in closed-ended vocal learners, song features crystallized early in life are less subject to changes in adulthood. This is why less attention has been paid to lifelong song changes in closed-ended learners. However, in the eyes of female birds that gain benefits from choosing mates based on male songs, not only past but also current conditions encoded in songs would be meaningful, given that even crystallized songs in closed-ended learners would not be identical in the long term. In this study, we examine within-individual song changes in the Java sparrow Lonchura oryzivora, with the aim of shedding light on the relationship between song and long-term life history. Specifically, we compared song length, tempo, and song complexity measures between the point just after song crystallization and around 1 yr later, and also compared those traits between fathers and sons to clarify the effect of vocal learning. While it is not surprising that song complexity did not differ depending on age or between fathers and sons, we found that song length and tempo increased with age. Follow-up analyses have revealed that frequency bandwidth and peak frequency of song notes also elevated with age. Our results show that song performance related to motor skills can be improved even after song crystallization. We also suggest that song performance in closed-ended vocal learners gives a reliable clue for mate choice by reflecting male quality with aging.
  • Hiroko Kagawa, Masayo Soma
    BEHAVIOURAL PROCESSES 99 138 - 144 0376-6357 2013/10 [Not refereed][Not invited]
     
    Bird songs have evolved under sexual selection pressure. Songs include multiple features that are subject to female preference, but recent comparative research has indicated evolutionary tradeoffs between song performance and complexity in some species. Trill, a repetition of the same sound, is a performance-related song trait; higher trill performance can be achieved at the cost of song complexity at the among-species or population level. The aim of this study was to examine whether such tradeoffs also account for within-species variation in Java sparrow songs, which include both multiple trill types and non-trill parts. We found a great individual variation in trill proportion, trill performance, and song complexity. A positive association between trill performance and body size suggested that trills can serve as an indicator of male quality. However, contrary to the tradeoffs predicted by previous studies based on other passerine species, trill performance and song complexity, i.e., note repertoire, were positively correlated: males in better condition can sing songs with larger note repertoires and higher trill performance, which may explain how trills and non-trill notes are both maintained and have co-evolved by sexual selection in Java sparrow songs. (C) 2013 Elsevier B.V. All rights reserved.
  • M. Soma, K. Okanoya
    BEHAVIOUR 150 (13) 1491 - 1508 0005-7959 2013 [Refereed][Not invited]
     
    Postmating sexual selection plays an important part in the evolution of secondary sexual characters. Based on differential allocation theory that predicts biased reproductive investment of females depending on the attractiveness of mates, a number of previous studies have shown that egg production is related in various ways to ornamental sexual traits of males, but evidence for behavioural sexual traits is less abundant. In this study we examine such maternal effects in relation to birdsong. Because the Bengalese finch is a monomorphic songbird, courtship song serves a key role in mate choice. To take into account individual female differences in egg production performance, we sequentially paired naive, captive, female Bengalese finches to two different males, and investigated if their reproductive investment (clutch size, egg mass and hatchling sex ratio) was related to the song traits of their mates. We found that clutch size and egg mass were highly repeatable within individual females while sex ratio was not. Despite the inflexibility of egg mass within each female, egg mass increased when females were mated to males with longer songs. In addition, we found a non-significant weak tendency toward male-biased sex ratio in relation to longer song duration of mates. Our findings suggest that females mated to better mates adjusted their reproductive investment by producing heavier eggs and possibly offspring of the more costly sex.
  • Masayo F. Soma
    ORNITHOLOGICAL SCIENCE 10 (2) 89 - 100 1347-0558 2011/12 [Not refereed][Not invited]
     
    Birdsong is an important sexually selected trait, and its acoustic features are socially transmitted in the process of song learning. Though a great deal of research has been conducted to shed light on the mechanisms and functions of song learning, we still do not have clear answers as to why for some species, including Estrildid finches and humans, the ability of "social" vocal learning has evolved. Therefore, this paper addresses social factors responsible for song learning, especially focusing on tutor choice in Estrildid finches, with the aim of elucidating what is already clear about how the social environment shapes songs and what is needed in future studies.
  • Ai Hasegawa, Masayo Soma, Toshikazu Hasegawa
    ORNITHOLOGICAL SCIENCE 10 (1) 73 - 80 1347-0558 2011/06 [Refereed][Not invited]
     
    Estrildine finches are important model species in experimental studies on female mating preferences, but research has focused on only two species from this family, namely, the Zebra Finch Taeniopygia guttata and the Bengalese Finch Lonchura striata var. domestica, and we know comparatively little about other closely related species. Therefore, we investigated sexual dimorphism and female choice in the Java Sparrow Padda oryzivora, which also belongs to the family Estrildidae, to open up the potential for comparative research and to better understand sexual selection in this family. First, we took measurements of eight morphological traits of six male and six female Java Sparrows: natural wing length, maximum wing length, tarsus length, tail length, exposed culmen length, bill width, bill depth and body mass. We quantitatively confirmed that Java Sparrows show sexual dimorphism in bill depth, with males having deeper bills than females. Second, we examined individual variation in male courtship songs by analyzing their acoustic and syntactical structure. After collecting data on male morphological and song-related traits, we conducted two-way choice tests to ask which kinds of male traits predicted which males were preferred by female Java Sparrows. In the choice tests, we put a cage with a female in between two cages each containing one male and recorded the position of the female every 30 seconds by point sampling. We performed a stepwise regression analysis to assess the relationship between the time females spent in front of each male and male morphological and song-related traits. The results indicated that females base their preference upon large body size, which is likely to act as a good indicator of male quality. However, no preference was observed for song-related traits or sexually dimorphic bill depth. Perhaps the sexual dimorphism in bill size is instead the evolutionary outcome of male-male competition.
  • Masayo Soma, Laszlo Zsolt Garamszegi
    BEHAVIORAL ECOLOGY 22 (2) 363 - 371 1045-2249 2011/03 [Not refereed][Not invited]
     
    The theory of sexual selection predicts a relationship between male sexual traits and reproductive success. This prediction has been tested extensively using the complexity of birdsong as a model for trait elaboration. However, contradictory results have emerged. Some studies have demonstrated that males with large repertoires enjoy a reproductive advantage, whereas other studies have failed to support this prediction. To make general inferences from this mixed evidence, we quantitatively reviewed the relevant literature using a meta-analytic approach. The mean effect size for the song/mating success association was significant, but the effects were generally weak, affected by publication bias, confounded by uncontrolled variables, and differing across the traits examined. Effect sizes were heterogeneous across studies due to species-specific effects, differences in mating systems, and song phenotypes. The degree of association between song complexity and reproductive success was independent of the strength of sexual selection, as assessed by the degree of polygyny and extrapair paternity. Our results highlight the importance of considering various biological factors to understand the role of repertoires in mediating mating success in different species.
  • Masayo Soma, Mariko Hiraiwa-Hasegawa, Kazuo Okanoya
    ANIMAL BEHAVIOUR 78 (5) 1107 - 1113 0003-3472 2009/11 [Not refereed][Not invited]
     
    Birdsong is an important sexually selected trait, and its acoustic features are socially transmitted in the process of song learning. The preference of some female songbirds for song complexity as measured by repertoire size has driven the evolution of many song traits and song-learning programs in males. If juvenile birds adopt the attractiveness of model birds by copying their repertoires, tutor choice in the early song-learning period should be crucial for future reproductive success. Thus, we hypothesized that the song quality of tutors influences the tutor choice of chicks in the Bengalese finch, Lonchura striata var. domestica. Each subject chick was reared by a foster pair in an individual breeding cage. At around the fledging period, we introduced another unrelated adult male (subtutor) into each cage to simulate the natural social environment in which chicks have opportunities to hear the songs of nonfather males after fledging. Subject chicks learned from both tutors when the total note repertoire size of the two models was small; they tended to learn from the subtutor when the father had a smaller note repertoire size. The acquired note repertoire size was not affected by whether subjects learned songs from the foster father or not. Song learning from the subtutor contributed to a larger note repertoire. Observed patterns in tutor choice, in particular song learning from a subtutor, could be an adaptive strategy that helps chicks compensate for exposure to smaller repertoire size in models. (C) 2009 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.
  • Masayo Soma, Mariko Hiraiwa-Hasegawa, Kazuo Okanoya
    BEHAVIORAL ECOLOGY AND SOCIOBIOLOGY 63 (3) 363 - 370 0340-5443 2009/01 [Not refereed][Not invited]
     
    Birdsong differs from other sexual traits in that the acquisition process involves learning. Especially in close-ended learning species like the Bengalese finch, conditions experienced during the critical song-learning period can have a profound influence on song quality. Therefore, to understand song evolution from a life-history perspective, we investigated early ontogenetic effects on song quality. In particular, we focused on maternal effects and sibling competition. In asynchronously hatching bird species, the age hierarchy among nestlings affects physical development due to competition for food; mothers may influence this competition by adjusting their investment in each egg according to its sequence in the laying order. To independently assess these effects, chicks of the Bengalese finch were cross-fostered so that the age hierarchies formed in fostered broods were independent of the laying order. Our results indicate that song quality partially reflects early ontogenetic conditions, whereas song duration and note-type repertoire were independent of either laying order or age hierarchy. The syntactical complexity of note order declined over the laying sequence. This finding suggests that the song learning ability is influenced by within-clutch variation in maternal investment toward eggs. Considering that song syntactical complexity is subject to female preference in the Bengalese finch, it is likely that maternal resource allocation strategies play a role in song evolution.
  • Masayo Soma, Toshikazu Hasegawa, Kazuo Okanoya
    JOURNAL OF AVIAN BIOLOGY 39 (1) 101 - 107 0908-8857 2008/01 [Not refereed][Not invited]
     
    Recent studies of the vocal mechanisms of songbirds have shown that there are physical effects on sound production. Interspecific and phylogenetic comparisons have shown that body mass and bill morphology are two major factors affecting vocalizations such as songs. We analyzed the distance calls of female Bengalese finches Lonchurastriata var. domestica, the non-vocal learning sex, to assess the potential physical effects on the acoustic structure of vocalizations. By experimentally controlling rearing condition using cross-fostering we could examine the effects of the developmental environment and genetic background. None of the tested factors affected the peak frequency of the distance calls, but we found that larger-billed birds tended to produce shorter bout calls with higher trill rates. These results suggest that the divergence of bill morphology can affect acoustic features at the within-population level. We also found that the birds reared in the same foster brood and siblings from the same genetic parents tended to produce calls with similar trill rates. This implies that the trill rate is under the influence of developmental and genetic factors.
  • Masayo Soma, Daichi S. Saito, Toshikazu Hasegawa, Kazuo Okanoya
    BEHAVIORAL ECOLOGY AND SOCIOBIOLOGY 61 (11) 1695 - 1705 0340-5443 2007/09 [Not refereed][Not invited]
     
    Maternal effects, such as investment in eggs, have profound effects on offspring fitness. Mothers are expected to skew their investment depending on the laying order and sex when unequal sibling competition occurs within a brood because of sex-specific vulnerability and age hierarchy caused by asynchronous hatching. The Bengalese finch hatches asynchronously and shows a moderate reversed sexual size dimorphism. However, contrary to commonly accepted assumptions of size-dependent vulnerability, the smaller sex (male) is more vulnerable to developmental stress caused by sibling competition. We investigated whether maternal investment would be biased by the position in laying order and the sex of eggs, and also explored the possible differences in growth patterns depending on sex, laying order, and age hierarchy by observing chicks fostered to experimentally manipulated broods where brood composition was controlled and age hierarchy was more enhanced than in natural breeding conditions. We found that overall patterns of maternal investment favored the disadvantageous sectors of sibling competition, i.e., eggs of later laying order and sons over those of early laying order and daughters. We also examined the effect of laying order on adult body size and sex differences in growth patterns. When reared in the subordinate age hierarchy, females could compensate for the deficit of decreased growth rate by taking longer to mature, whereas males could not. We suggest that this sex-specific growth pattern could be the cause of sex differences in vulnerability to early developmental stress.
  • Masayo Soma, Miki Takahasi, Maki Ikebuchi, Hiroko Yamada, Madoka Suzuki, Toshikazu Hasegawa, Kazuo Okanoya
    ETHOLOGY 112 (11) 1071 - 1078 0179-1613 2006/11 [Not refereed][Not invited]
     
    Birdsong is an acoustic ornament. According to indicator models, a trait must be costly to act as an honest signal, but the potential costs of elaborate songs are still poorly understood. The developmental stress hypothesis Suggests that learned song characteristics Could be an honest indicator of early developmental conditions because the brain structures associated with learning songs are susceptible to early developmental stress, which could thus affect song development. Unlike previous studies of developmental stress that examined the effect of a stress hormone or restricted nutrition, we observed Bengalese finches under semi-natural breeding conditions in captivity to investigate the relationship between early rearing conditions (e.g., brood size and sex ratio) and the subsequent variation in body size and song among individuals. Our results suggest that the early rearing environment directly affects body size and song complexity, whereas song output is determined mainly by' body size. These results support the developmental stress hypothesis. Moreover, Our findings are the first to show that developmental condition affects not only the number of note types but also the syntactical complexity of the song.
  • M Soma, T Hasegawa
    BEHAVIOUR 141 1121 - 1134 0005-7959 2004/09 [Not refereed][Not invited]
     
    Social foraging is a far more complex phenomenon than individual foraging because of the many social interactions and communications that affect individual behaviour. Social dominance and social learning markedly influence the foraging efficiency of individuals in an unfamiliar environment. This study investigated how these two factors affect the costs and benefits of social foraging. We presented a novel feeding environment to captive budgerigars, Melopsittacus undulatus, and then compared the latency to feeding and number of pecks at the food for social and individual conditions, and for high-, middle-, and low-ranking birds. When in the social condition, birds started foraging faster and pecked more food than in the individual condition. Presumably, this lowered neophobia and enhanced foraging efficiency in the social condition was caused by social facilitation. Low-ranking birds had less resource accessibility in the social condition, probably because they were constrained by the existence of higher-ranking birds when it came to accessing the feeder. Nevertheless, the food intake of low-ranking birds almost equaled that of high- or middle-ranking birds in the social condition. In summary, high status is definitely an advantage, while low status adds some costs to individuals. Nevertheless, low-ranking birds compensate for this through enhanced foraging. It was clear that social foraging provides a great advantage to foragers of each rank, because of social facilitation.

Awards & Honors

  • 2016 資生堂女性研究者サイエンスグラント
     
    受賞者: 相馬雅代
  • 2015 文部科学大臣表彰 若手科学者賞
     
    受賞者: 相馬雅代
  • 2011 日本鳥学会 黒田賞
     
    受賞者: 相馬雅代
  • 2005 第30 回鳥類内分泌研究会若手奨励賞
     
    受賞者: 相馬雅代

Educational Activities

Teaching Experience

  • Behavioral Control System Sciences
    開講年度 : 2018
    課程区分 : 修士課程
    開講学部 : 生命科学院
    キーワード : 行動,脳,中枢神経系,社会性,コミュニケーション,認知,神経回路,ニューロン,最初期遺伝子,感覚情報処理,運動制御,学習,進化
  • Inter-Graduate School Classes(General Subject):Natural and Applied Sciences
    開講年度 : 2018
    課程区分 : 修士課程
    開講学部 : 大学院共通科目
    キーワード : 生物多様性、遺伝的多様性、陸上植物、土壌微生物、藻類、哺乳類、鳥類、魚類、昆虫、陸水生物、海洋動物 Biodiversity, genetic variation, terrestrial plants, soil microorganisms, algae, mammals, birds, fishes, insects, freshwater organisms, marine animals
  • Fundamental Lecture in Biological Diversity
    開講年度 : 2018
    課程区分 : 修士課程
    開講学部 : 環境科学院
    キーワード : 生物多様性、遺伝的多様性、陸上植物、土壌微生物、藻類、哺乳類、鳥類、魚類、昆虫、陸水生物、海洋動物 Biodiversity, genetic variation, terrestrial plants, soil microorganisms, algae, mammals, birds, fishes, insects, freshwater organisms, marine animals
  • Inter-Graduate School Classes(General Subject):Natural and Applied Sciences
    開講年度 : 2018
    課程区分 : 修士課程
    開講学部 : 大学院共通科目
    キーワード : 生命システム, 生命機能, 研究方法論, 研究技術論
  • Biosystems Science
    開講年度 : 2018
    課程区分 : 修士課程
    開講学部 : 生命科学院
    キーワード : 生命システム, 生命機能, 研究方法論, 研究技術論
  • Biostatistics
    開講年度 : 2018
    課程区分 : 学士課程
    開講学部 : 理学部
    キーワード : 統計,データ解析
  • Biology II
    開講年度 : 2018
    課程区分 : 学士課程
    開講学部 : 全学教育
    キーワード : 生物の多様性,系統,進化,生物の形態,生命活動の多様性
  • Laboratory Course in Behavioral Neurobiology
    開講年度 : 2018
    課程区分 : 学士課程
    開講学部 : 理学部
    キーワード : 行動、脳、中枢神経系、脊椎動物、無脊椎動物、組織化学、遺伝子発現、光学計測、社会行動、コミュニケーション
  • Brain, Behavior and Evolution
    開講年度 : 2018
    課程区分 : 学士課程
    開講学部 : 理学部
    キーワード : 動物行動、進化、コミュニケーション、微小脳、視覚、嗅覚、学習、認知


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