Time flies: A bibliometric exploration of evolving trends in Drosophila -based research

Abstract

Drosophila melanogaster has been used as a model organism in biomedical research for over a century, facilitating a number of fundamental breakthroughs and Nobel Prize-winning discoveries in genetics, developmental biology and disease mechanisms. This bibliometric analysis assessed 140,962 Drosophila-related publications indexed in the Web of Science Core Collection from 1984 to 2024. Publication output increased rapidly until the late 1990s, with an average annual growth rate of 13%, before stabilising. Co-authorship analysis revealed extensive international collaborative networks spanning multiple continents, while keyword co-occurrence analysis identified seven major research clusters: developmental biology, cell biology, circadian biology, ageing, evolutionary biology, molecular biology and immunology. Temporal trend analyses demonstrated sustained growth in fields such as neuroscience, neurodegenerative diseases, ageing, circadian biology, immunology, pharmacology, toxicology and environmental sciences. These findings provide a comprehensive overview of contemporary Drosophila-based research, highlighting its ongoing global relevance, evolving applications and emerging research frontiers. The unique advantages of Drosophila for investigating complex biological processes establish it as a more ethical and efficient alternative to vertebrate models in modern biomedical science.

Introduction

Drosophila is a genus of flies belonging to the family Drosophilidae. Drosophila melanogaster, commonly known as the fruit fly or vinegar fly, is a quintessential model organism in biomedical research, particularly in studies that leverage genetics as a primary strategy.^1,2 The Drosophila genome comprises approximately 14,000 protein-coding genes distributed across four pairs of chromosomes. ³ Despite its relative simplicity compared to the human genome, roughly 75% of human disease genes have functional homologues in Drosophila. ⁴ For over a century, this model has provided profound insights into fundamental biological processes and has contributed to several Nobel Prize-winning discoveries. ⁵ Notable laureates include: Morgan (1933) for chromosomal heredity; Müller (1948) for X-ray-induced mutations; Lewis, Nüsslein-Volhard and Wieschaus (1995) for embryonic development; Hoffmann (2011) for innate immunity; and Hall, Rosbash and Young (2017) for circadian rhythms (as well as Axel (2004) for the olfactory system, although the work was primarily based on rodent studies).

Beyond this notable scientific heritage, Drosophila serves as a pivotal experimental system that bridges the gap between unicellular organisms, such as yeast, and more complex vertebrates like mice and zebrafish. ⁶ Its utility is driven by several advantageous features, ⁷ including high fecundity, large progeny numbers and short generation intervals, which facilitate large-scale or high-throughput experimentation with low maintenance space and costs. These traits have also led to the widespread global use of Drosophila in biomedical innovation projects in resource-limited regions, through initiatives such as DrosAfrica and Droso4Nigeria. ⁸ Furthermore, the organism’s relatively small genome and ease of genetic manipulation facilitate the use of cutting-edge research strategies and technologies.

Decades of intensive study have yielded a robust ecosystem of fly-specific resources, such as FlyBase, while the evolutionary conservation of key pathways contributes to the acceleration of human medical advances. Finally, research based on the use of Drosophila presents fewer ethical constraints compared to the use of vertebrate models, making it an appropriate tool for both advanced research and educational outreach, as demonstrated by the Manchester Fly Facility.^9,10 The ease of fly husbandry and the depth of biological understanding render Drosophila a convenient organism for use in the classroom, with a range of curriculum-relevant topics and micro-experiments that can be undertaken in schools.

Bibliometric analysis has emerged as a rigorous methodology for quantitatively evaluating research landscapes and forecasting academic trends. ¹¹ Although several conventional narrative-type reviews of Drosophila-based research exist,^12–14 the current bibliometric analysis aims to perform the literature review from an analytical, rather than a descriptive, perspective. In general, bibliometric approaches aim to identify key patterns and relationships, reveal conceptual structures and research gaps, direct the path of future research, and inform policy development. ¹⁵ Despite the organism’s significance, such analyses remain scarce in the Drosophila literature, with existing reports often limited to specific subfields like cancer research. ¹⁶ The primary objective of this analysis was to unveil critical themes and trace the evolution of research paradigms within the broader Drosophila field. By providing a panoramic view of contemporary trends and collaborations, this work serves as a visual resource to guide future discoveries and identify emerging research frontiers.

Methods

To ensure data consistency, a search of the Web of Science Core Collection was conducted in July 2025, retrieving documents published from 1 January 1984 through to 31 December 2024. The search strategy employed the topic tag ‘TS=(Drosophila)’, with the scope limited to original research articles and reviews. Other publication types, such as meeting abstracts, book chapters, letters and editorials, were excluded. Document metadata, including authors, affiliations, countries, journal title, publication year, keywords, subject areas and citation topics, were extracted and stored in CSV format.

The Web of Science platform assigns journals to specific subject areas. Journals categorised under ‘Multidisciplinary Sciences’ or ‘General and Internal Medicine’ were reclassified more precisely, as appropriate. Citation topics were derived from algorithmic citation clusters. While meso-topics were manually labelled based on thematic content, micro-topics were algorithmically assigned by using the most significant keywords. A single document may be indexed under multiple research areas or citation topics.

Bibliometric networks were constructed and visualised using VOSviewer (version 1.6.20; Leiden University, the Netherlands). International collaborative connectivity was mapped through co-authorship analysis based on jointly authored publications. In these visualisations, node size corresponds to a country’s total publication output, with larger nodes indicating greater output. The lines (links) represent collaborative ties. Keyword co-occurrence networks were generated by calculating the frequency of shared keywords, which were organised into thematic clusters to identify research patterns within the literature. ¹⁷ To assess longitudinal trends, the annual publication count served as a proxy for global research interest. Data trends were modelled using polynomial regression smoothing splines plotted against the publication year. ¹⁸ The interpretation of these bibliometric findings followed the ‘3Ss’ framework of sensemaking: scanning, sensing and substantiating. ¹⁹

Results

A total of 140,962 Drosophila-related publications were identified in the Web of Science Core Collection between 1984 and 2024. The vast majority were published in English (n = 139,566; 99%), followed by Russian (n = 828), French (n = 273), Chinese (n = 76) and Spanish (n = 61). Original articles accounted for approximately 90% of the retrieved publications (n = 126,993), with reviews comprising the remaining 13,969 documents. Publication output increased rapidly from 1984 to 1998, with an average annual growth rate of 13% (Figure 1). Following this period, productivity stabilised, reaching a peak in 2013 with 4765 publications (4290 articles and 475 reviews).

Figure 1.

Annual trends in the number of research publications based on the use of Drosophila (1984–2024).

The top five contributors to Drosophila-based research were the USA (n = 67,222), Germany (n = 12,966), UK (n = 12,021), China (n = 10,387) and Japan (n = 10,345). While Japan followed a trajectory similar to that of Western nations, characterised by initial growth followed by a plateau, Russian publications began to emerge in the post-Soviet era before stabilising (Figure 2). Publication output from China began to accelerate in the mid-1990s following economic reforms. By 2024, China ranked second globally in academic productivity. As shown in Figure 3, the co-authorship network displayed intricate international collaborations. These co-authorship clusters consistently included countries across multiple continents, underscoring the field’s global nature. Leading institutions are listed in Table 1, featuring large organisations such as the University of California System (Research Organization Registry ID 00pjdza24) and the Max Planck Society (Research Organization Registry ID 01hhn8329). It should be noted that the institutions ranked at the top of the table are primarily parent organisations. For example, the University of London (Research Organization Registry ID 04cw6st05) encompasses several member institutions, including King’s College London, Queen Mary University of London, UCL (University College London) and the Institute of Cancer Research, London. While the highest-ranked institutions are predominantly in the USA and Western Europe, the Russian Academy of Sciences, the Chinese Academy of Sciences and the University of Tokyo also demonstrate significant output.

Figure 2.

Fitted polynomial regression smoothing splines illustrating publication trends in selected countries for research based on the use of Drosophila (1984–2024).

Figure 3.

International co-authorship network in research based on the use of Drosophila (1984–2024).

Table 1.

Top 20 institutions by Drosophila research output in terms of publications (1984–2024).

Rank	Lead author affiliation	No. of publications retrieved
1	University of California System, USA	9694
2	Centre national de la recherche scientifique (CNRS), France	6705
3	Howard Hughes Medical Institute, USA	6386
4	Harvard University, USA	4553
5	Max Planck Society, Germany	2962
6	National Institutes of Health (NIH), USA	2853
7	Russian Academy of Sciences, Russia	2756
8	University of Cambridge, UK	2620
9	Institut National de la Santé et de la Recherche Médicale (INSERM), France	2500
10	University of Texas System, USA	2449
11	University of London, UK	2102
12	University of California Berkeley, USA	2058
13	Consejo Superior de Investigaciones Cientificas (CSIC), Spain	1956
14	Stanford University, USA	1956
15	Chinese Academy of Sciences, China	1880
16	Cornell University, USA	1824
17	University of Tokyo, Japan	1792
18	University System of Ohio, USA	1757
19	Universite Paris-Saclay, France	1734
20	University of California San Francisco, USA	1700

The top journals, according to the number of published Drosophila-based research articles during the study period, were Development (n = 4438), Proceedings of the National Academy of Sciences of the United States of America (n = 4213), Genetics (n = 4063), Journal of Biological Chemistry (n = 3233), PLoS One (n = 3198) and Developmental Biology (n = 2706). Table 2 ranks the key areas and citation themes based on the total number of retrieved publications, identifying molecular biology, genetics and developmental biology as the core pillars of the field. Granular micro-topic analysis highlighted prominent subfields, such as epigenetic regulation, transposable elements, olfactory systems, ageing mechanisms and circadian rhythms.

Table 2.

Top 20 subject areas and featured topics in Drosophila research (1984–2024).

Rank	Research area and number of publications retrieved		Meso-topics and number of relevant publications identified		Micro-topics and number of relevant publications identified
1	Biochemistry/molecular biology	39,359	Molecular & cell biology: Cancer & development	29,187	Drosophila development	22,455
2	Genetics/heredity	29,256	Entomology	20,448	Insect cold adaptation	5988
3	Cell biology	26,378	Molecular & cell biology: Genetics	15,881	Juvenile hormone	5649
4	Developmental biology	15,739	Cell biology	6832	Epigenetic regulation	5619
5	Science technology: Other topics	14,761	Molecular & cell biology: Cancer, autophagy & apoptosis	5715	Sexual selection	4571
6	Life sciences/biomedicine: Other topics	10,096	Zoology & animal ecology	4765	Transposable elements	4292
7	Neurosciences/neurology	9643	Phylogenetics & genomics	4250	Olfactory systems	3986
8	Evolutionary biology	9107	Smell & taste science	4065	Ageing mechanisms	3724
9	Entomology	8990	Longevity	3859	Neural crest	3502
10	Environmental sciences/ecology	7336	Sleep science & circadian biology	3281	Invertebrate immunity	3357
11	Zoology	6519	Neurodegenerative diseases	3169	Circadian-melatonin	3120
12	Biotechnology/applied microbiology	4609	Molecular & cell biology: DNA damage	2609	Microtubule dynamics	3106
13	Physiology	4537	Micro & long noncoding RNA	2528	Alternative splicing	2543
14	Biophysics	3831	Molecular & cell biology: Physiology	2376	Genetic diversity	1777
15	Oncology	2297	Animal sensing	2166	Membrane trafficking	1728
16	Chemistry	2260	Neuroscience	2059	Insect sensory systems	1656
17	Immunology	1949	Chromosome disorders	1572	Insect symbiosis	1547
18	Pharmacology/pharmacy	1885	Crop science	1207	Evolutionary dynamics	1544
19	Toxicology	1841	Molecular & cell biology: Mitochondria	1167	RNAi	1435
20	Research/experimental medicine	1720	Genome studies	1154	Heat shock proteins	1391

Keyword co-occurrence analysis identified seven major thematic clusters (Figure 4). The red cluster (developmental biology) included keywords such as ‘pattern’, ‘induction’ and ‘cell fate’. The blue cluster (cell biology) featured keywords like ‘growth’, ‘apoptosis’ and ‘cancer’. The orange cluster (circadian biology) contained keywords such as ‘rhythms’, ‘light’ and ‘period’. The purple cluster (ageing biology) included keywords like ‘oxidative stress’, ‘lifespan’ and ‘longevity’. The yellow cluster (evolutionary biology) contained keywords such as ‘evolution’, ‘selection’ and ‘fitness’. The green cluster (molecular biology) included keywords like ‘chromatin’, ‘interference’ and ‘DNA methylation’. Finally, the cyan cluster (immunology) featured keywords such as ‘recognition’, ‘Drosophila Toll’ and ‘NF-kappa B’. The density of connections between these clusters reflects the high degree of interdisciplinary synergy in the field.

Figure 4.

Thematic keyword co-occurrence map for research based on the use of Drosophila (1984–2024).

Longitudinal analysis revealed shifting trends across subject areas (Figure 5). Classical disciplines, including Biochemistry/Molecular Biology, Genetics/Heredity, Cell Biology, Developmental Biology, and Evolutionary Biology, showed significant early growth followed by recent marginal declines. Interestingly, while cancer research with Drosophila (Figure 5, Oncology) experienced an early surge, the number of publications has since moderated. Conversely, steady growth was observed in the fields of Neuroscience, Immunology, and Toxicology. Notably, publications in Pharmacology/Pharmacy and Environmental Sciences/Ecology have consistently risen over the past four decades.

Figure 5.

Publication trends in featured subject areas of research based on the use of Drosophila (1984–2024), fitted with polynomial regression smoothing splines.

Among curated micro- and meso-topics, early academic publication growth was driven by classical subjects such as epigenetic regulation, sexual selection and alternative splicing. However, similar to transposable elements, these topics have recently seen a decline (Figure 6). In contrast, research into ageing mechanisms, circadian biology, olfactory systems, invertebrate immunity and neurodegenerative diseases exhibits sustained or increasing momentum, perhaps representing the current frontiers of research based on the use of Drosophila.

Figure 6.

Publication trends in featured micro-topics and meso-topics of research based on the use of Drosophila (1984–2024), fitted with polynomial regression smoothing splines.

Discussion

In the current era of exponential growth in the number of scientific publications, bibliometric analysis has become a valuable tool for understanding the dynamics of a specific research field. This study elucidates the rise and evolution of research based on the use of Drosophila over the past 40 years (i.e. between 1984 and 2024, inclusive). Modern biology has relied heavily on Drosophila-based studies, which have provided fundamental insights into a range of biological processes, that have been subsequently extrapolated to other organisms. The rapidly increasing number of such publications observed in this study, from 1984 up until the early 2010s, marks this era as a ‘golden age’ for Drosophila as a valuable resource in biomedical research. Early breakthroughs, such as gene cloning and hybridisation, led to a boom in the discovery of fundamental biological mechanisms. Today, sophisticated technologies like the GAL4/UAS system, RNA interference and CRISPR-Cas9 facilitate precise genetic manipulation, with CRISPR-Cas9 enabling the introduction of human disease mutations into orthologous Drosophila genes. ²⁰ Hence, it is anticipated that Drosophila models will continue to be used in human disease research across multiple areas. In the current analysis, fluctuations in the number of publications relating to a certain field of study served as a proxy for trends in academic output within that field.

Trends within specific fields of study

Oncology

A previous study by Kamdem et al. ¹⁶ showed a continuous growth in the number of publications on ‘Drosophila cancer models’ between 2008 and 2018. However, the present study revealed an ongoing decline in oncology-related published literature from 2008. This discrepancy likely arises from our analysis being limited to publications classified under the subject area of ‘Oncology’, potentially overlooking related research categorised in areas such as Biochemistry/Molecular Biology or Cell Biology. Thus, the trends observed in this study may be influenced by the publication classification process. Although flies do not naturally develop cancer in the same way that mammals do, the substantial overlap between oncogenic transformation and developmental processes make Drosophila-based studies useful for identifying oncogenes and tumour suppressors. ²¹ Signalling pathways such as Notch, Hippo and BMP, which are dysregulated in many cancers, were first characterised in Drosophila. ²² As a whole-animal model for studying tumour biology, cancer genetics and cancer drug discovery, Drosophila represents a cutting-edge system whose experimental versatility complements mammalian models in uncovering metastatic mechanisms, tumour–microenvironment interactions, and immune-related aspects of cancer research.

Neuroscience

Drosophila-based research has led to some significant advances in the field of neuroscience.^23,24 Several milestones in neurobiology, such as the discovery of potassium and TRP channels, circadian rhythms, sexual behaviour and olfactory communication, have been established by using Drosophila. ²⁵ Recently, neurodegenerative diseases, characterised by the progressive loss of neuronal structure and function, have become prominent areas of study using these models. ²⁶ In the case of Alzheimer’s disease, the pathological effects of amyloid-β and tau protein accumulation have been extensively examined. For Parkinson’s disease, relevant publications have steadily increased since the introduction of a viable model in 2000. ²⁷ Additionally, emerging studies are investigating the aetiology and pathophysiology of Huntington’s disease. Despite the challenges of assessing behavioural and cognitive aspects in flies, as well as the differences in pathophysiology between ‘knock-in’ experiments and the slow, progressive nature of the human diseases, it is likely that Drosophila-based research will continue to shed light on neurodegenerative mechanisms and help in identifying therapeutic opportunities.

Immunology and infectious diseases

Like other insects, Drosophila are constantly exposed to environmental pathogens, primarily bacteria. The innate immune system of Drosophila shares many similarities with the human immune system, which plays a central role in defence against microbial infections. ²⁸ The observed decrease in the number of immunology-based publications may be attributed to the clear distinction between human adaptive immunity and the antimicrobial peptides of Drosophila. Consequently, Drosophila is not a suitable model for studying antibody- and lymphocyte-dependent adaptive immune responses. However, this limitation can be potentially advantageous, as it results in a stronger focus on innate immune responses, particularly innate immune memory. ²⁹

Ageing mechanisms

The publication trend for this type of study over the analysed period showed an initial rise, followed by stabilisation. The short lifespan of the fruit fly permits the examination of gene effects in older individuals, while studying ageing mechanisms in mammals is constrained by their much longer lifespans. Drosophila have been used to explore the interactions between mutation accumulation and antagonistic pleiotropy in the evolution of ageing, as well as the trade-offs between longevity and other fitness traits, such as nutritional and metabolic factors. ³⁰ However, as noted in an editorial by Clancy et al., ⁵ the use of Caenorhabditis elegans has surpassed that of Drosophila in ageing studies, due to advantages in frozen storage and an even shorter lifespan. Nonetheless, it is important to recognise that the short lifespan of these invertebrate models may not accurately reflect long-term chronic diseases in humans. Close communication and collaboration across diverse disciplines can facilitate the effective implementation of cross-species model organism approaches, and serve to uncover the fundamental biological underpinnings of health and disease. ³¹

Pharmacology/pharmacy and toxicology

The number of publications in these two research areas has been steadily increasing over the years. While cell-based screening is the most widely used method for identifying potential bioactive compounds in drug discovery, the results often translate poorly to in vivo testing, as toxicity profiles in cell cultures do not accurately reflect tissue-specific responses. ¹⁶ Compared to expensive, time-consuming rodent models, tailored screening platforms employing Drosophila are particularly appealing due to their low husbandry costs, high reproductive rates and reduced ethical concerns. Substances that negatively affect development or physiological processes can be quickly eliminated, and off-target effects can be systematically detected. ³² In this context, Drosophila can also be utilised for toxicological studies from early development and into adulthood. These studies typically involve exposing flies to various compounds — such as pesticides, organic solvents and metals — and assessing their effects on survival, reproduction, development and behaviour. ³³ Consequently, there is a growing trend to further leverage Drosophila for use in pharmacological and toxicological screening.

Future prospects

Overall, the intermediate complexity of Drosophila provides a valuable whole-organism context for scientific discovery, encompassing development, behaviour and physiology. Drosophila offers several advantages over vertebrates, including speed, cost-effectiveness, high fecundity, fewer ethical considerations and less need for specialised infrastructure. As such, Drosophila can effectively replace or significantly reduce the use of vertebrates as a ‘first-line’ in vivo system. This helps support the Three Rs principles ³⁴ (replacement, reduction and refinement) relating to vertebrate animal use, and is in alignment with ethical and sustainability goals that prioritise low resource consumption while maintaining scientific rigour and translational relevance. However, limitations on the use of Drosophila remain in areas such as cardiovascular research and other chronic mammalian diseases, which often require hybrid approaches.

Our bibliometric analysis demonstrated sustained global research output and robust international collaboration rooted in a transformative tradition of adapting classical methods to modern uses. This indicates that Drosophila remains a cornerstone of biological research, rather than it being a declining legacy model. Looking ahead, Drosophila will continue to serve as a resource-sustainable and ethically favourable model, generating further impact across multiple research areas. These areas could include the accelerated functional elucidation of gene sets and genetic variants, support for system-level analyses of complex biological processes, and expedited in vivo screening in pharmacology and toxicology. Rising publication trends highlight several emerging and expanding domains, particularly in neuroscience and neurodegenerative diseases, ageing mechanisms and lifespan regulation, circadian biology, invertebrate immunity and environmental sciences. These fields could benefit from the unique experimental advantages of Drosophila and may present promising opportunities for systems biology, multi-omics integration and in silico–in vivo based approaches.

Limitations of the current study

Several limitations of this study should be acknowledged. Our data were exclusively derived from the Web of Science Core Collection. Although the Web of Science is a multidisciplinary and authoritative citation index database, the omission of other databases, such as Scopus, may result in variations in the quantity of documents and other metrics. Additionally, using Drosophila as the search topic inevitably included publications unrelated to the model organism, such as research in the fields of entomology and infectious diseases. Furthermore, the document quality was not assessed as part of this study, and (as is the case in all bibliometric analyses) the data presented are subject to potential publication bias, where positive or novel findings are more likely to be published. While bibliometric analysis is inherently quantitative and objective, it still requires interpretation, which introduces an element of subjectivity. ¹⁹ Due to space limitations, a detailed discussion of some representative studies was not possible. However, despite these constraints, bibliometric research presents a valuable opportunity to contribute to theory and practice. ³⁵

Conclusions

In the current atmosphere of funding shortages that demotivate investigators and, more importantly, impact the long-term career choices of many young scientists, ³⁶ this study reflects the substantial efforts devoted to Drosophila-based research over the past four decades. It provides a comprehensive temporal, thematic and structural perspective on Drosophila-based research. The analysis draws attention to the fact that Drosophila offers a useful approach to addressing the complexities of biological research, as well as helping to accelerate discovery and facilitate innovative developments.

The bibliometric analysis serves to highlight the past contribution of this model organism to our understanding of biology, as well as the evolving applications of its use in research. Serving as the foundational organism for classical genetics during the first half of the 20th century, and then as a driving force in developmental biology during the latter half, Drosophila exemplifies how related approaches have since expanded across an ever-growing range of biological fields, facilitated by the many advantages of fly-based research. While its utility in classical fields like developmental and evolutionary biology persists, Drosophila-based research is carving out a contemporary niche and being used to explore research frontiers in areas such as neuroscience, immunology, ageing studies, pharmacology and toxicology.