Morphogenesis and form  ·  Article

Synthetic morphology: modular morphogenetic events

Synthetic morphology is the effort to build form in living cells by switching on single morphogenetic events through engineered gene expression. Davies and Cachat describe a set of modules that control proliferation, elective cell death, cell fusion, cell adhesion and locomotion.3 The aim is an artificial developmental cycle that runs from patterning through gene expression to shape.2 The term itself is older than the modules.1

Earliest held
1911, Leduc, S.
Most discussed in
Horizontal mitochondrial transfer as a novel…, 2024
In the library
27 passages in 18 works
Rewritten
2026-10-22
01

An older phrase

The phrase appears in Leduc's 1911 book The Mechanism of Life. Leduc reports a memoir of 1871 by the Dutch scholar Harting on the artificial production of some organic calcareous formations. Citing Professor R. Dubois, he says the memoir had cost Harting more than thirty years of work.1 Dubois called synthetic morphology a field still in its infancy. He hoped that, given as much time as had passed since the first artificial production of urea, it would progress as far as its older sister, synthetic chemistry.1 In this early sense the term meant making organic forms artificially, not engineering cells.

02

Ten events and the modules

Davies and Cachat (2016) trace the modern programme to an early speculative paper on synthetic morphogenesis. That paper suggested that most mammalian morphogenesis combines about ten basic events: proliferation, cell death, fusion, adhesion, de-adhesion, migration, two transitions between epithelium and mesenchyme, epithelial folding and lumen formation.2 They add that earlier studies had found genes, some of viral origin, whose activation drives one specific event. This allowed their group to build modules for proliferation, elective cell death, cell fusion, cell adhesion and locomotion.3

03

Coupling to patterning

Davies and Cachat state that the modules were published separately from work on patterning. In principle they can be placed downstream of it.3 A simple proof-of-principle example has been made. It used patterning by phase separation, then triggered a cell death module in one colour of cells. This turned a pattern of colours into a net-like arrangement of cells and spaces.3 Davies and Glykofrydis (2020) say the modules were first shown only in very simple verification assays. They describe the same sieve-like result and report that work is underway to couple patterning to responses more sophisticated than elimination.4

04

Fusion and later uses

Cell fusion is also studied outside the module programme. Kippenberger and colleagues (2023) note that Physarum can transfer a learned behaviour to other individuals by cell fusion. They take this to indicate a material correlate of learning, for which they say there is no convincing theory.7 Reid (2023) disputes Vogel and Dussutour's (2016) description of unhabituated plasmodia acquiring a learned behaviour from habituated ones. Reid argues that after fusion, the regions should not be called individual plasmodia.6 Iorio and colleagues (2024) report that fusing human mesenchymal stem cells with cardiomyocytes moves mitochondria into the cardiomyocytes and reprograms the hybrids toward a progenitor-like state.8 Abramson and Levin (2021) offer behaviorist methods to the field of functional synthetic morphology.5 Pai and colleagues (2025) treat synthetic morphology as a source of model systems free of a history of environmental selection.9

SourcesEach quotation was checked word for word against the passage it opens.
  1. Synthetic morphology is yet only in its infancy, let us hope that in a time equal to that which has already expired since the first artificial production of ureaLeduc, S., 1911 · The Mechanism of Life · open at passage 290
  2. suggested that most examples of mammalian morphogenesis use combinations of approximately ten basic morphogenetic events.Davies JA, Cachat E, 2016 · Synthetic biology meets tissue engineering · open at passage 13
  3. This observation has allowed us to produce a set of modules for ‘synthetic morphology’, that allow control of proliferation, elective cell death, cell fusion, cell adhesion and locomotion [5].Davies JA, Cachat E, 2016 · Synthetic biology meets tissue engineering · open at passage 14
  4. patterning by phase separation generates patches of one phase amongst cells of the other phase, then a cell death programme is induced in the patches to leave a sieve-like sheet of cells with holesDavies JA, Glykofrydis F, 2020 · Engineering pattern formation and morphogenesis · open at passage 14
  5. We provide an overview of the conceptual and methodological tools that classical behaviorism has to offer the field of functional synthetic morphologyAbramson CI, Levin M, 2021 · Behaviorist approaches to investigating memory and learning: A primer for… · open at passage 9
  6. it is incorrect to label these regions as individual plasmodia after cell–cell fusion has occurred.Reid CR, 2023 · Thoughts from the forest floor: a review of cognition in the slime mould… · open at passage 41
  7. Of note, this work shows that Physarum can transfer the learned behavior to other individuals by cell fusion.Kippenberger S, Pipa G, Steinhorst K, Zöller N, Kleemann J…, 2023 · Learning in the Single-Cell Organism Physarum polycephalum: Effect of Propofol · open at passage 2
  8. Cell fusion between human MSCs and cardiomyocytes results in MT into cardiomyocytes, and the resulting hybrid cells are reprogrammed toward a progenitor-like state [150].Iorio R, Petricca S, Mattei V, Delle Monache S, 2024 · Horizontal mitochondrial transfer as a novel bioenergetic tool for mesenchymal… · open at passage 39
  9. Tremendous progress in the fields of synthetic biology, synthetic morphology, and biorobotics has made available biological model systems that are unencumbered by a history of environmental selection forcesPai VP, Pio-Lopez L, Sperry MM, Erickson P, Tayyebi P…, 2025 · Basal Xenobot transcriptomics reveals changes and novel control modality in… · open at passage 29
Linked ideas
The general article on the field. This one covers the specific modular morphogenetic events.
Davies and Cachat place the modules within a developmental cycle for building tissues.
Constructs of this kind are the model systems that Pai and colleagues describe as products of synthetic morphology.
Concerns constructing novel living forms, the broader aim behind engineering morphology.
Physarum learning transferred by cell fusion, and the dispute over what counts as one individual.
Basal cognitionrelated to
Abramson and Levin pair synthetic morphology with behaviorist tests of memory and learning in novel organisms.
Evolvabilityrelated to
Davies & Glykofrydis list evolvability and robustness as goals for elaborating engineered patterns, while expanding the morphogenetic module toolbox. The passages never directly link evolvability to the cell death, fusion or adhesion modules.
Where it is discussedPassages matching synthetic morphology, morphogenetic modules, cell adhesion module, elective cell death
2024Horizontal mitochondrial transfer as a novel bioenergetic tool for mesenchymal… · Iorio R, Petricca S, Mattei V, Delle…4
2016Synthetic biology meets tissue engineering · Davies JA, Cachat E3
2021Behaviorist approaches to investigating memory and learning: A primer for… · Abramson CI, Levin M3
2025Basal Xenobot transcriptomics reveals changes and novel control modality in… · Pai VP, Pio-Lopez L, Sperry MM…2
2020Engineering pattern formation and morphogenesis · Davies JA, Glykofrydis F2
2024Embryos assist morphogenesis of others through calcium and ATP signaling… · Tung A, Sperry MM, Clawson W, Pavuluri…1
2023Thoughts from the forest floor: a review of cognition in the slime mould… · Reid CR1
2025Aging as a Loss of Goal-Directedness: An Evolutionary Simulation and Analysis… · Pio-Lopez L, Hartl B, Levin M1
2026Mind Everywhere: A Framework for Conceptualizing Goal-Directedness in Biology… · Levin M, Resnik DB1
2022Technological Approach to Mind Everywhere: An Experimentally-Grounded Framework… · Levin M1