Fly Development

From Embryology
Revision as of 11:11, 30 April 2020 by Z8600021 (talk | contribs)
(diff) ← Older revision | Latest revision (diff) | Newer revision → (diff)
Embryology - 14 Jun 2024    Facebook link Pinterest link Twitter link  Expand to Translate  
Google Translate - select your language from the list shown below (this will open a new external page)

العربية | català | 中文 | 中國傳統的 | français | Deutsche | עִברִית | हिंदी | bahasa Indonesia | italiano | 日本語 | 한국어 | မြန်မာ | Pilipino | Polskie | português | ਪੰਜਾਬੀ ਦੇ | Română | русский | Español | Swahili | Svensk | ไทย | Türkçe | اردو | ייִדיש | Tiếng Việt    These external translations are automated and may not be accurate. (More? About Translations)


Fly animation.gif
Drosophila melanogaster drawing

This page introduces the fly, drosophila, as a developmental model organism. The small drosophila fruitfly has been used by genetisists for many years now and much is now understood about its development in relation to gene expression and regulatory mechanisms.

In recent years, using developmental mutants, many mechanisms of development in the fly have been shown to be almost identical to those seen in humans and other animals. In fact, these developmental mechanisms have become the "paradigm" for our understanding of development.

The fruitfly (drosophila) was and is the traditional geneticist's tool. It has been transformed to an magnificent tool for the embryologist, with many developmental mechanisms being uncovered in this system combined with homolgy gene searches in other species.

Drosophila researchers have received to date received 5 Nobel prizes (1933, 1995, 2011). The most recent in 2011 "for their discoveries concerning the activation of innate immunity".

There is also a difference in basic body structure between males and females, males lack the seventh abdominal segment (A7) present in females. This has recently been shown to be due to a down-regulation of epidermal growth factor receptor (EGFR) activity and fewer histoblasts in the male A7 in the early pupae.[1]

Fly Links: ANAT2341 Project (2009) | Homeobox | Category:Fly

Some Recent Findings

Drosophila gastrulation
Drosophila gastrulation[2]
Scanning EM of adult fly head
  • The Physical Mechanisms of Drosophila gastrulation: mesoderm and endoderm Invagination[2] "A critical juncture in early development is the partitioning of cells that will adopt different fates into three germ layers: the ectoderm, the mesoderm, and the endoderm. This step is achieved through the internalization of specified cells from the outermost surface layer, through a process called gastrulation. In Drosophila, gastrulation is achieved through cell shape changes (i.e., apical constriction) that change tissue curvature and lead to the folding of a surface epithelium. Folding of embryonic tissue results in mesoderm and endoderm invagination, not as individual cells, but as collective tissue units. The tractability of Drosophila as a model system is best exemplified by how much we know about Drosophila gastrulation, from the signals that pattern the embryo to the molecular components that generate force, and how these components are organized to promote cell and tissue shape changes. For mesoderm invagination, graded signaling by the morphogen, Spätzle, sets up a gradient in transcriptional activity that leads to the expression of a secreted ligand (Folded gastrulation) and a transmembrane protein (T48). Together with the GPCR Mist, which is expressed in the mesoderm, and the GPCR Smog, which is expressed uniformly, these signals activate heterotrimeric G-protein and small Rho-family G-protein signaling to promote apical contractility and changes in cell and tissue shape. A notable feature of this signaling pathway is its intricate organization in both space and time. At the cellular level, signaling components and the cytoskeleton exhibit striking polarity, not only along the apical-basal cell axis, but also within the apical domain. Furthermore, gene expression controls a highly choreographed chain of events, the dynamics of which are critical for primordium invagination; it does not simply throw the cytoskeletal "on" switch." fly
  • Multimodal transcriptional control of pattern formation in embryonic development[3] "Predicting how interactions between transcription factors and regulatory DNA sequence dictate rates of transcription and, ultimately, drive developmental outcomes remains an open challenge in physical biology. Using stripe 2 of the even-skipped gene in Drosophila embryos as a case study, we dissect the regulatory forces underpinning a key step along the developmental decision-making cascade: the generation of cytoplasmic mRNA patterns via the control of transcription in individual cells. Using live imaging and computational approaches, we found that the transcriptional burst frequency is modulated across the stripe to control the mRNA production rate. However, we discovered that bursting alone cannot quantitatively recapitulate the formation of the stripe and that control of the window of time over which each nucleus transcribes even-skipped plays a critical role in stripe formation. Theoretical modeling revealed that these regulatory strategies (bursting and the time window) respond in different ways to input transcription factor concentrations, suggesting that the stripe is shaped by the interplay of 2 distinct underlying molecular processes."
  • N-cadherin orchestrates self-organization of neurons within a columnar unit in the Drosophila medulla[4] "The columnar structure is a basic unit of the brain, but its developmental mechanism remains unknown. The medulla, the largest ganglion of the fly visual center, provides a unique opportunity to reveal the mechanisms of three-dimensional organization of the columns. We reveal that column formation is initiated by three core neurons that establish distinct concentric domains within a column. We demonstrate the in vivo evidence of N-cadherin-dependent differential adhesion among the core columnar neurons within a column along a two-dimensional layer in the larval medulla. The two-dimensional larval columns evolve to form three distinct layers in the pupal medulla. We propose the presence of mutual interactions among the three layers during formation of the three-dimensional structures of the medulla columns." neural
More recent papers  
Mark Hill.jpg
PubMed logo.gif

This table allows an automated computer search of the external PubMed database using the listed "Search term" text link.

  • This search now requires a manual link as the original PubMed extension has been disabled.
  • The displayed list of references do not reflect any editorial selection of material based on content or relevance.
  • References also appear on this list based upon the date of the actual page viewing.

References listed on the rest of the content page and the associated discussion page (listed under the publication year sub-headings) do include some editorial selection based upon both relevance and availability.

More? References | Discussion Page | Journal Searches | 2019 References | 2020 References

Search term: Drosophila Development

Older papers  
These papers originally appeared in the Some Recent Findings table, but as that list grew in length have now been shuffled down to this collapsible table.

See also the Discussion Page for other references listed by year and References on this current page.

  • AnnoFly: Annotating Drosophila Embryonic Images Based on an Attention-Enhanced RNN Model[5] "The Berkeley Drosophila Genome Project (BDGP) has collected a large-scale spatial gene expression database for studying Drosophila embryogenesis. Given the expression images, how to annotate them for the study of Drosophila embryonic development is the next urgent task. In order to speed up the labor-intensive labeling work, automatic tools are highly desired. ...To address these challenges, we develop a new annotator for the fruit fly embryonic images, called AnnoFly. Driven by an attention-enhanced RNN model, it can weight images of different qualities, so as to focus on the most informative image patterns. We assess the new model on three standard data sets. The experimental results reveal that the attention-based model provides a transparent approach for identifying the important images for labeling, and it substantially enhances the accuracy compared with the existing annotation methods, including both single-instance and multi-instance learning methods."AnnoFly
  • High throughput in vivo functional validation of candidate congenital heart disease genes in Drosophila[6] "We developed a Drosophila-based functional system to screen candidate disease genes identified from Congenital Heart Disease (CHD) patients. 134 genes were tested in the Drosophila heart using RNAi-based gene silencing. Quantitative analyses of multiple cardiac phenotypes demonstrated essential structural, functional, and developmental roles for more than 70 genes, including a subgroup encoding histone H3K4 modifying proteins. We also demonstrated the use of Drosophila to evaluate cardiac phenotypes resulting from specific, patient-derived alleles of candidate disease genes. We describe the first high throughput in vivo validation system to screen candidate disease genes identified from patients." cardiovascular abnormalities
  • Lineage-associated tracts defining the anatomy of the drosophila first instar larval brain[7] "Fixed lineages derived from unique, genetically specified neuroblasts form the anatomical building blocks of the Drosophila brain. Neurons belonging to the same lineage project their axons in a common tract, which is labeled by neuronal markers. In this paper, we present a detailed atlas of the lineage-associated tracts forming the brain of the early Drosophila larva, based on the use of global markers (anti-Neuroglian, anti-Neurotactin, Inscuteable-Gal4>UAS-chRFP-Tub) and lineage-specific reporters. We describe 68 discrete fiber bundles that contain axons of one lineage or pairs/small sets of adjacent lineages. Bundles enter the neuropil at invariant locations, the lineage tract entry portals. Within the neuropil, these fiber bundles form larger fascicles that can be classified, by their main orientation, into longitudinal, transverse, and vertical (ascending/descending) fascicles. We present 3D digital models of lineage tract entry portals and neuropil fascicles, set into relationship to commonly used, easily recognizable reference structures such as the mushroom body, the antennal lobe, the optic lobe, and the Fasciclin II-positive fiber bundles that connect the brain and ventral nerve cord. Correspondences and differences between early larval tract anatomy and the previously described late larval and adult lineage patterns are highlighted."
  • Development of the imaginal wing disc[8] "LIM-HD gene tailup (islet), together with the HD genes of the iroquois complex, specify the notum territory of the disc. Later, tailup has been shown to act as a prepattern gene that antagonizes formation of sensory bristles on the notum of this fly. ...We conclude that tailup acts on bristle development by several, even antagonistic, mechanisms."
  • The four-dimensional pattern of fly neuron development[9] "We show that segment-specific generation of the Ap cluster neurons is achieved by the integration of the anteroposterior and temporal cues in several different ways. Generation of the Ap neurons in abdominal segments is prevented by anteroposterior cues stopping the cell cycle in the stem cell at an early stage. In brain segments, late-born neurons are generated, but are differently specified due to the presence of different anteroposterior and temporal cues. Finally, in thoracic segments, the temporal and spatial cues integrate on a highly limited set of target genes to specify the Ap cluster neurons."
  • Heart Development[10] "We used an optical coherence tomography imaging technique that provided images similar to echocardiography in humans to measure the cardiac function in adult flies. We identified mutants in members of the rhomboid protease family and epidermal growth factor receptor that cause an enlarged cardiac chamber. Interestingly, abnormalities in the function of members of the epidermal growth factor receptor family in humans that undergo certain chemotherapies are associated with the development of dilated cardiomyopathy and heart failure. Our results suggest that epidermal growth factor receptor signaling may be an evolutionarily conserved pathway that is necessary to maintain normal adult cardiac function."


melanogaster group

Taxonomy Id: 32346 Rank: species group

Genetic code: Translation table 1 (Standard) Mitochondrial genetic code: Translation table 5 Lineage( abbreviated ): Eukaryota; Metazoa; Arthropoda; Tracheata; Hexapoda; Insecta; Pterygota; Neoptera; Endopterygota; Diptera; Brachycera; Muscomorpha; Ephydroidea; Drosophilidae; Drosophila


The drosophila lifespan varies with temperature and is about 30 days at 29 °C.

A series of papers published between 1976 to 1979 by Turner and Mahowald, characterised the stages of drosophila development in beautiful scanning electron microscope (SEM) images.[11][12][13]

Hox Genes

Fly wild-type head.jpg Fly antennapedia head.jpg
Fly wild-type head[14] Fly antennapedia mutant head[14]

This is the classic mutation that gave rise to the discovery of Hox genes and other genes related to body pattern formation. In this mutant during development the fly embryo incorrectly positioned where (antenna) should have be two legs (pedia)[14]. The discovery of this mutant in Walter Gehring's lab opened up the field of developmental genes and this field has been rewarded with the 1995 Nobel prize in Medicine.

Links: Hox | 1995 Nobel Prize

Hippo Genes

The Hippo (Hpo) pathway, first identified in Drosophila, controls organ size by regulating cell proliferation (inhibition) and apoptosis (induction). In contrast, the TOR signalling pathway regulates organ size by stimulating cell growth, thus increasing cell size.

Fly Phenotype (dorsal view head thorax SEM)
Fly Hippo-type dorsal view head thorax SEM.jpg Fly WT dorsal view head thorax SEM.jpg
Hippo-type (hpo) Wild-type (WT)
Image source[15]
Links: Developmental Signals - Hippo

Neural Development

Summary of neural development from neural stem cell population and the gene regulation involved.[9]

Fly neural development 01.png


  1. Foronda D, Martín P & Sánchez-Herrero E. (2012). Drosophila Hox and sex-determination genes control segment elimination through EGFR and extramacrochetae activity. PLoS Genet. , 8, e1002874. PMID: 22912593 DOI.
  2. 2.0 2.1 Martin AC. (2020). The Physical Mechanisms of Drosophila Gastrulation: Mesoderm and Endoderm Invagination. Genetics , 214, 543-560. PMID: 32132154 DOI.
  3. Lammers NC, Galstyan V, Reimer A, Medin SA, Wiggins CH & Garcia HG. (2020). Multimodal transcriptional control of pattern formation in embryonic development. Proc. Natl. Acad. Sci. U.S.A. , 117, 836-847. PMID: 31882445 DOI.
  4. Trush O, Liu C, Han X, Nakai Y, Takayama R, Murakawa H, Carrillo JA, Takechi H, Hakeda-Suzuki S, Suzuki T & Sato M. (2019). N-cadherin orchestrates self-organization of neurons within a columnar unit in the Drosophila medulla. J. Neurosci. , , . PMID: 31175213 DOI.
  5. Yang Y, Zhou M, Fang Q & Shen HB. (2019). AnnoFly: Annotating Drosophila Embryonic Images Based on an Attention-Enhanced RNN Model. Bioinformatics , , . PMID: 30601935 DOI.
  6. Zhu JY, Fu Y, Nettleton M, Richman A & Han Z. (2017). High throughput in vivo functional validation of candidate congenital heart disease genes inDrosophila. Elife , 6, . PMID: 28084990 DOI.
  7. Hartenstein V, Younossi-Hartenstein A, Lovick JK, Kong A, Omoto JJ, Ngo KT & Viktorin G. (2015). Lineage-associated tracts defining the anatomy of the Drosophila first instar larval brain. Dev. Biol. , 406, 14-39. PMID: 26141956 DOI.
  8. de Navascués J & Modolell J. (2010). The pronotum LIM-HD gene tailup is both a positive and a negative regulator of the proneural genes achaete and scute of Drosophila. Mech. Dev. , 127, 393-406. PMID: 20580820 DOI.
  9. 9.0 9.1 Karlsson D, Baumgardt M & Thor S. (2010). Segment-specific neuronal subtype specification by the integration of anteroposterior and temporal cues. PLoS Biol. , 8, e1000368. PMID: 20485487 DOI.
  10. Yu L, Lee T, Lin N & Wolf MJ. (2010). Affecting Rhomboid-3 function causes a dilated heart in adult Drosophila. PLoS Genet. , 6, e1000969. PMID: 20523889 DOI.
  11. Turner FR & Mahowald AP. (1976). Scanning electron microscopy of Drosophila embryogenesis. 1. The structure of the egg envelopes and the formation of the cellular blastoderm. Dev. Biol. , 50, 95-108. PMID: 817949
  12. Turner FR & Mahowald AP. (1977). Scanning electron microscopy of Drosophila melanogaster embryogenesis. II. Gastrulation and segmentation. Dev. Biol. , 57, 403-16. PMID: 406152
  13. Alper PR. (1975). Letter: Lawsuit motivation. J Leg Med (N Y) , 3, 7. PMID: 1081572
  14. 14.0 14.1 14.2 Turner FR & Mahowald AP. (1979). Scanning electron microscopy of Drosophila melanogaster embryogenesis. III. Formation of the head and caudal segments. Dev. Biol. , 68, 96-109. PMID: 108157
  15. Johnson R & Halder G. (2014). The two faces of Hippo: targeting the Hippo pathway for regenerative medicine and cancer treatment. Nat Rev Drug Discov , 13, 63-79. PMID: 24336504 DOI.


Developmental Dynamics

Journal of Neurobiology

Online Textbooks

Molecular Biology of the Cell (4th Edn) Alberts, Bruce; Johnson, Alexander; Lewis, Julian; Raff, Martin; Roberts, Keith; Walter, Peter. New York: Garland Publishing; 2002.

Developmental Biology (6th Edn) Gilbert, Scott F. Sunderland (MA): Sinauer Associates, Inc.; c2000.

Search NLM Online Textbooks- "drosophila development" : Molecular Biology of the Cell | Molecular Cell Biology | The Cell- A molecular Approach


Sen A & Cox RT. (2017). Fly Models of Human Diseases: Drosophila as a Model for Understanding Human Mitochondrial Mutations and Disease. Curr. Top. Dev. Biol. , 121, 1-27. PMID: 28057297 DOI.

Tadros W & Lipshitz HD. (2005). Setting the stage for development: mRNA translation and stability during oocyte maturation and egg activation in Drosophila. Dev. Dyn. , 232, 593-608. PMID: 15704150 DOI.

Pereanu W & Hartenstein V. (2004). Digital three-dimensional models of Drosophila development. Curr. Opin. Genet. Dev. , 14, 382-91. PMID: 15261654 DOI.

Voas MG & Rebay I. (2004). Signal integration during development: insights from the Drosophila eye. Dev. Dyn. , 229, 162-75. PMID: 14699588 DOI.

Weigmann K, Klapper R, Strasser T, Rickert C, Technau G, Jäckle H, Janning W & Klämbt C. (2003). FlyMove--a new way to look at development of Drosophila. Trends Genet. , 19, 310-1. PMID: 12801722 DOI.


Shingleton AW, Das J, Vinicius L & Stern DL. (2005). The temporal requirements for insulin signaling during development in Drosophila. PLoS Biol. , 3, e289. PMID: 16086608 DOI.

Search PubMed

Search Aug2005 "drosophila development" 13228 reference articles of which 1899 were reviews.

Search Pubmed: fly development | drosophila development

External Links

External Links Notice - The dynamic nature of the internet may mean that some of these listed links may no longer function. If the link no longer works search the web with the link text or name. Links to any external commercial sites are provided for information purposes only and should never be considered an endorsement. UNSW Embryology is provided as an educational resource with no clinical information or commercial affiliation.


There are a number of excellent internet resources for Fly development.

Fly Pages

Animal Development: axolotl | bat | cat | chicken | cow | dog | dolphin | echidna | fly | frog | goat | grasshopper | guinea pig | hamster | horse | kangaroo | koala | lizard | medaka | mouse | opossum | pig | platypus | rabbit | rat | salamander | sea squirt | sea urchin | sheep | worm | zebrafish | life cycles | development timetable | development models | K12
Historic Embryology  
1897 Pig | 1900 Chicken | 1901 Lungfish | 1904 Sand Lizard | 1905 Rabbit | 1906 Deer | 1907 Tarsiers | 1908 Human | 1909 Northern Lapwing | 1909 South American and African Lungfish | 1910 Salamander | 1951 Frog | Embryology History | Historic Disclaimer

Glossary Links

Glossary: A | B | C | D | E | F | G | H | I | J | K | L | M | N | O | P | Q | R | S | T | U | V | W | X | Y | Z | Numbers | Symbols | Term Link

Cite this page: Hill, M.A. (2024, June 14) Embryology Fly Development. Retrieved from

What Links Here?
© Dr Mark Hill 2024, UNSW Embryology ISBN: 978 0 7334 2609 4 - UNSW CRICOS Provider Code No. 00098G