Secreted signaling molecules act as morphogens to control patterning and growth in many developing tissues. Since locally produced morphogens spread to form a concentration gradient in the surrounding tissue, spreading is generally thought to be the key step in the non‐autonomous actions. Here, we review recent advances in tool development to investigate morphogen function using the role of decapentaplegic (Dpp)/bone morphogenetic protein (BMP)‐type ligand in the Drosophila wing disc as an example. By applying protein binder tools to (...) distinguish between the roles of Dpp spreading and local Dpp signaling, we found that Dpp signaling in the source cells is important for wing patterning and growth but Dpp spreading from this source cells is not as strictly required as previously thought. Given recent studies showing unexpected requirements of long‐range action of different morphogens, manipulating endogenous morphogen gradients by synthetic protein binder tools could shed more light on how morphogens act in developing tissues. (shrink)

The veins are cuticular structures that differentiate in precise patterns in insect wings. The genetic and molecular basis of vein pattern formation in Drosophila melanogaster is beginning to be unravelled with the identification and characterisation of the gene products that position the veins and direct their differentiation. Genes affecting the veins fall into two groups: transcriptional regulators that specify individual veins, and members of signalling pathways involved in patterning and differentiation of the veins. The elaboration of the vein pattern (...) is progressive in time and requires the coordinated activities of these signalling pathways and the transcription factors regulated by them. Although the network of genetic interactions that determine vein cell fate is well understood, very little is known about the cellular biology underlying the acquisition of vein histotype. BioEssays 25:443–451, 2003. © 2003 Wiley Periodicals, Inc. (shrink)

The imaginal discs of Drosophila melanogaster, which form the adult epidermal structures, are a good experimental model for studying morphogenesis. The genital disc forms the terminalia, which are the most sexually dimorphic structures of the fly. Both sexes of Drosophila have a single genital disc formed by three primordia. The female genital primordium is derived from 8th abdominal segment and is located anteriorly, the anal primordium (10 and 11th abdominal segments) is located posteriorly, and the male (...) genital primordium from the 9th abdominal segment lies between them. In both sexes, only two of these three primordia develop to form the adult terminalia. The anal primordium develops in both sexes but, depending on the genetic sex, will form either male or female analia. However, only one of the genital primordia develops in each sex, forming either the male or the female genitalia. This depends on the genetic sex of the fly. Therefore, the genital disc is a very good experimental model of how the sex‐determination and homeotic genes — which determine cell identity — interact to direct the development of a population of cells into male or female terminalia. It has been proposed that the sexually dimorphic development of the genital disc is the result of an integrated genetic input, made up by the sex‐determination gene doublesex and the homeotic gene Abdominal‐B. This input acts by modulating the response to Hedgehog, Wingless, and Decapentaplegic morphogenetic signals. BioEssays 23:698–707, 2001. © 2001 John Wiley & Sons, Inc. (shrink)

Drosophila imaginal discs (appendage primordia) have proved invaluable for deciphering cellular and molecular mechanisms of animal development. By combining the accessibility of the discs with the genetic tractability of the fruit fly, researchers have discovered key mechanisms of growth control, pattern formation and long‐range signaling. One of the principal experimental attractions of discs is their anatomical simplicity — they have long been considered to be cellular monolayers. During larval stages, however, the growing discs are 2‐sided sacs composed of a (...) columnar epithelium on one side and a squamous ‘peripodial’ epithelium on the other. Recent studies suggest important roles for peripodial epithelia in processes previously assumed to be confined to columnar cell monolayers. BioEssays 23:691–697, 2001. © 2001 John Wiley & Sons, Inc. (shrink)

Many animals display a capacity to regenerate tissues or even a complete body. One of the main goals of regenerative biology is to identify the genes and genetic networks necessary for this process. Drosophila offers an ideal model system for such studies. The wide range of genetic and genomic approaches available for use in flies has helped in initiating the deciphering of the mechanisms underlying regeneration, and the results may be applicable to other organisms, including mammals. Moreover, most models (...) of regeneration require experimental manipulation, whereas in Drosophila discrete domains can be ablated by genetically induced methods. Here, we present a summary of current research into imaginal disc regeneration and discuss the power of this tissue as a tool for understanding the genetics of regeneration. (shrink)

The particular fascination of butterfly wings for developmental biologists (and others) lies in their spectacular array of colour patterns. The evolutionary and developmental relationships between these patterns have been analysed and we know something of the cell interactions involved in their formation(1). Now butterfly homologues of Drosophila wing‐patterning genes have been identified, and their expression patterns offer the first clues to the molecular mechanisms which specify wing colour patterns(2).

In the past few years it has become apparent that the anterior/posterior (A/P) and dorsal/ventral (D/V) compartmant boundaries serve as the source of longrange signals that organize the A/P and D/V axes of the Drosophila wing. Recent work suggests that the vestigial gene may function as a nodal point through which the growth‐controlling activity of these two patterning systems is integrated(1).

The link between oncogenesis and normal development is well illustrated by the study of the Wnt family of proteins. The first Wnt gene (int‐1) was identified over a decade ago as a proto‐oncogene, activated in response to proviral insertion of a mouse mammary tumor virus. Subsequently, the discovery that Drosophila wingless, a developmentally important gene, is homologous to int‐1 supported the notion that int‐1 may have a role in normal development. In the last few years it has been recognized (...) that int‐1 and Wingless belong to a large family of related glyco‐proteins found in vertebrates and invertebrates. In recognition of this, members of this family have been renamed Wnts, an amalgam of int and Wingless. Investigation of Wnt genes in Xenopus and mouse indicates that Wnts have a role in cell proliferation, differentiation and body axis formation. Further analysis in Drosophila has revealed that Wingless function is required in several developmental processes in the embryo and imaginal discs. In addition, a genetic approach has identified some of the molecules required for the transmission and reception of the Wingless signal. We will review recent data which have contributed to our growing understanding of the function and mechanism of Drosophila Wingless signaling in cell fate determination, growth and specification of pattern. (shrink)

The Drosophila position‐specific (PS) integrins are members of the integrin family of cell surface receptors and are thought to be receptors for extracellular matrix components. Each PS integrin consists of an α subunit, αPS1 or αPS2, and a βPS subunit. Mutations in the βPS subunit and the αPS2 subunit have been characterised and reveal that the PS integrins have an essential role in the adhesion of different cell layers to each other. The PS integrins are especially required for the (...) function of the cell‐matrix‐cell junctions, where the muscles attach to the epidermis and where one surface of the developing wing adheres to the other. These junctions are similar to vertebrate focal adhesions and hemidesmosomes, which also contain integrins. Integrin‐mediated cell to cell adhesion via the extracellular matrix provides a way for tissues to adhere to each other without intermingling of their cells. (shrink)

Cells in multicellular organisms often do not intermingle freely with each other. Differential cell affinities can contribute to organizing cells into different tissues. Drosophila limbs and the vertebrate central nervous system are subdivided into compartments. Cells in adjacent compartments do not mix. Cell interactions mediated by Notch-family receptors have been implicated in the specification of these compartment boundaries. Two recent reports analyze the role of the Notch signaling pathway in the generation of an affinity boundary in the Drosophila (...) wing.1,2 The first report1 analyzes the connection between Notch and the actin cytoskeleton. The second report2 analyzes the differential requirements of Notch and the transcription factor Supressor of Hairless in generating the affinity boundary. BioEssays 28: 113–116, 2006. © 2006 Wiley periodicals, Inc. (shrink)

The Transforming growth factor beta (TGF‐β) family of secreted proteins regulates a variety of key events in normal development and physiology. In mammals, this family, represented by 33 ligands, including TGF‐β, activins, nodal, bone morphogenetic proteins (BMPs), and growth and differentiation factors (GDFs), regulate biological processes as diverse as cell proliferation, differentiation, apoptosis, metabolism, homeostasis, immune response, wound repair, and endocrine functions. In Drosophila, only 7 members of this family are present, with 4 TGF‐β/BMP and 3 TGF‐β/activin ligands. Studies (...) in the fly have illustrated the role of TGF‐β/BMP ligands during embryogenesis and organ patterning, while the TGF‐β/activin ligands have been implicated in the control of wing growth and neuronal functions. In this review, we focus on the emerging roles of Drosophila TGF‐β/activins in inter‐organ communication via long‐distance regulation, especially in systemic lipid and carbohydrate homeostasis, and discuss findings relevant to metabolic diseases in humans. -/- . (shrink)

The appendages of Drosophila develop from the imaginal discs. During the extensive growth of these discs cell lineages are for the most part unfixed, suggesting a strong role for cell‐cell interactions in controlling the final pattern of differentiation. However, during early and middle stages of development, discs are subdivided by strict lineage restrictions into a small number of spatially distinct compartments. These compartments appear to be maintained by stably inheriting states of gene expression; the compartmentspecific expression of two such (...) ‘selector’ ‐ like genes, engrailed and apterous, are critical for anterior‐posterior and dorso‐ventral compartmentalization, respectively. Recent work suggests that one purpose of compartmentalization is to establish regions of specialized cells near compartment boundaries via intercompartmental induction, using molecules like the hedgehog protein. Thus, compartments can act as organizing centers for patterning within compartments. Evidence for non‐compartmental patterning mechanisms will also be discussed. (shrink)

The Drosophila compound eye is an excellent experimental system for analysing fate induction of identifiable single cells. Each ommatidium, a unit eye, contains eight photoreceptors (R1‐R8), and the differentiation of these photoreceptors occurs in the larval eye imaginal disc in discrete steps: first R8 is determined, then R2/R5, R3/R4, R1/R6 and finally R7. Induction of R7, in particular, has been extensively studied at the molecular level. The R8 photoreceptor presents on its surface a ligand, Bride of Sevenless, that (...) binds and activates Sevenless receptor tyrosine kinase in the R7 precursor. Autophosphorylated Sevenless initiates a Ras1‐mediated cascade, which eventually activates transcription factors in the nucleus via Raf1 and MAP kinases, resulting in R7 development. However, recent studies indicate that Sevenless (Sev) functions just to neuralize the cell and has no role in R7 fate determination per se. It appears that the R7 fate may represent the lowest rung of a ‘neuronal ground state’, which is attained without any specific inductive cue. It is plausible that the R7 precursor is actively prevented from taking on the neuronal fate and this inhibition is removed by activation of Sev. (shrink)

The cuticular surface of Drosophila is decorated by parallel arrays of polarized structures such as hairs and sensory bristles; for example, on the wing each cell produces a distally pointing hair. These patterns are termed [tissue polarity]. Several genes are known whose activity is essential for the development of normal tissue polarity. Mutations in these genes alter the orientation of the hair or bristle with respect to neighboring cells and the body as a whole. The phenotypes of mutations (...) in these genes allows them to be placed in three phenotypic groups. Based on their behavior in genetic mosaics, it has proved possible to determine that individual genes are required either for the generation of an intercellular polarity signal and/or the transduction of that signal to the cytoskeleton. (shrink)

Determination of cell fate in the developing eye of Drosophila depends on a precise sequence of cellular interactions which generate the stereotypic array of ommatidia. In the eye imaginal disc, an initially unpatterned epithelial sheath of cells, the first step in this process may be the specification of R8 photoreceptor cells at regular intervals. Genes such as Notch and scabrous, known to be involved in bristle development, alos participate in this process, suggesting that the specification of ommatidial founder (...) cells and the formation of sensory organs in the adult epidermis may involve a similar mechanism, that of lateral inhibition. The subsequent steps of ommatidial assembly, following R8 assignment, involve a different mechanism: Undetermined cells read their position based on the contacts they make with neighbors that have already begun to differentiate. The development of the R7 photoreceptor cell, one of the eight photoreceptor cells in the ommatidium, is best understood. An important role seems to be played by sevenless, a receptor tyrosine kinase on the surface of the R7 precursor. It transmits the positional information– most likely encoded by the boss protein on the neighboring R8 cell membrane – into the cell via its tyrosine kinase, which activates a signal transduction cascade. Constitutive activation of the sevenless kinase by overexpression of an N‐terminally truncated form results in the diversion of other ommatidial cells into the R7 pathway suggesting that activation of the sevenless signalling pathway is sufficient to specify R7 development. Genetic dissection of this pathway should therefore identify components of a signalling cascade activated by a tyrosine kinase. (shrink)

What initiates vertebrate limb development and induces limbs to form where they do? For several years the answer to this intriguing question has been framed in terms of a working model that limb induction depends on a dialogue between two members of the Fibroblast Growth Factor (FGF) family of intercellular signaling molecules, FGF8 and FGF10. Now, a recent paper has written roles for signals encoded by WNT genes, the vertebrate relatives of the Drosophila wingless gene, into the script.(1) BioEssays (...) 23:865–868, 2001. © 2001 John Wiley & Sons, Inc. (shrink)

Molecular and developmental studies of limb pattern formation have recently gained widespread attention. The fact that vertebrate limbs are amenable to both genetic and embryological manipulations has established this model system as a valuable paradigm for studying vertebrate development. Limb buds are polarised along all three major axes and the establishment of the dorso‐ventral (DV) polarity is dependent upon cues localised in the trunk, where a DV ectodermal interface is produced by confrontation of dorsal and ventral identities. By analogy to (...) Drosophila imaginal disc development, this interface has been proposed to determine and position an ectodermal organising centre, the Apical Ectodermal Ridge (AER), controlling limb bud outgrowth. Recent fate mapping studies(1) and studies of genes regulating DV limb polarity(2‐6), AER formation(7,8) and differentiation(9) suggest, however, that DV patterning and AER induction, though coordinately regulated during limb bud outgrowth, may early on be more dissociated than expected. (shrink)

Planar cell polarity (PCP)‐signaling and associated tissue polarization are evolutionarily conserved. A well documented feature of PCP‐signaling in vertebrates is its link to centriole/cilia positioning, although the relationship of PCP and ciliogenesis is still debated. A recent report in Drosophila established that Frizzled (Fz)‐PCP core signaling has an instructive input to polarized centriole positioning in non‐ciliated Drosophila wing epithelia as a PCP read‐out. Here, we review the impact of this observation in the context of recent descriptions of (...) the relationship(s) of core Fz‐PCP signaling and cilia/centriole positioning in epithelial and non‐epithelial cells. All existing data are consistent with a model where Fz‐PCP signaling functions upstream of centriole/cilia positioning, independent of ciliogenesis. The combined data sets indicate that the Fz‐Dsh PCP complex is instructive for centriole/ciliary positioning via an actin‐based mechanism. Thereby, centriole/cilia/centrosome positioning can be considered an evolutionarily conserved readout and common downstream effect of PCP‐signaling from flies to mammals. (shrink)

Dlg (Discs large), Scrib (Scribble) and Lgl (Lethal giant larvae) are evolutionarily conserved components of a common genetic pathway that link the seemingly disparate functions of cell polarity and cell proliferation in epithelial cells. dlg, scrib and lgl have been identified as tumour suppressor genes in Drosophila, mutations of which cause similar phenotypes, involving disruption of cell polarity and neoplastic overgrowth of tissues. The molecular mechanisms by which Dlg, Scrib and Lgl proteins regulate cell proliferation are not clear, but (...) there is some evidence that epithelial polarisation is required for this regulation. Dlg, Scrib and Lgl are highly conserved between human and Drosophila, and we discuss evidence that these proteins also play a role in cancer progression in humans. BioEssays 25:542–553, 2003. © 2003 Wiley Periodicals, Inc. (shrink)

Recent studies of gene expression in the developing fruitfly leg support a model – Meinhardt's Boundary Model – which seems to contradict the prevailing paradigm for pattern formation in the imaginal discs of Drosophila – the Polar Coordinate Model. Reasoning from geometric first principles, this article examines the strengths and weaknesses of these hypotheses, plus some baffling phenomena that neither model can comfortably explain. The deeper question at issue is: how does the fly's genome encode the three‐dimensional anatomy of (...) the adult? Does it demarcate territories and boundaries (as in a geopolitical map) and then use those boundaries and their points of intersection as a scaffolding on which to erect the anatomy (the Boundary Model)? Or does it assign cellular fates within a relatively seamless coordinate system (the Polar Coordinate Model)? The existence of hybrid Cartesian‐polar models shows that the alternatives may not be so clear‐cut: a single organ might utilize different systems that are spatially superimposed or temporally sequential. (shrink)

Sensory organs are specialized to receive different kinds of input from the outside world. However, common features of their development suggest that they could have a shared evolutionary origin. In a recent paper, Niwa et al.1 show that three Drosophila adult sensory organs all rely on the spatial signals Decapentaplegic and Wingless to specify their position, and the temporal signal ecdysone to initiate their development. The proneural gene atonal is an important site for integration of these regulatory inputs. These (...) results suggest the existence of a primitive sensory organ precursor, which would differentiate according to the identity of its segment of origin. The authors argue that the eyeless gene controls eye disc identity, indirectly producing an eye from the sensory organ precursor within this disc. BioEssays 26:825–828, 2004. © 2004 Wiley Periodicals, Inc. (shrink)

In this paper we describe a test for Nijhout's hypothesis that the eyespot patterns on butterfly wings are the result of a threshold reaction of the epidermal cells to a concentration gradient of a diffusing degradable morphogen produced by focal cells at the centre of the future eyespot. The wings of the nymphalid butterfly, Bicyclus anynana, have a series of eyespots, each composed of a white pupil, a black disc and a gold outer ring. In earlier extirpation and transplantation (...) experiments it has been established that these eyespots are indeed organised around groups of signalling cells active during the first hours of pupal development. If these cells were to supply the positional information for eyespot formation in accordance with Nijhout's diffusion-degradation gradient model, then, when two foci are close together, the signals should sum, and this effect should be apparent in the detailed shape of the resulting pigment pattern. We give an equation for the form of the contours that would be obtained in this manner. We use this to test the morphogen gradient hypothesis on measurements of the outlines of fused eyespots obtained either by grafting focal cells close together, or by using a mutation that produces adjacent fused eyespots. The contours of the fused patterns were found to satisfy our equation, thus corroborating Nijhout's hypothesis to the extent possible with this particular type of experiment. (shrink)

Insect vision is nothing if not active. The regular head movements, called saccades, enable the fly Drosophila to keep a straight path in flight despite inequalities in the thrust of the wings. Using their own motion, bees in flight measure the ranges of nearby objects. A long history of research shows that bees discriminate visually in ways that depend on their activity or task, so we must distinguish between vision during flying, fixating or hovering and landing.Bees return again and (...) again for a reward of sugar solution and use their eyes to find their way. In an apparatus that makes them discriminate between two simulataneously visible but regularly interchanged targets, seen at a distance of 27 cm, bees are able to distinguish a remarkable number of simple patterns, but they fail in certain critical cases. The results can be explained with the hypothesis that bees have several broadly tuned overlapping filters with large fields that respond to the predominant orientation in a region of the image, and others for radial and circular patterns. Together with colour, these filters are independent of range. Bees prefer to use landmarks where they can, then global pattern at the largest scale, and lastly the detail around the goal. The way that discrimination of one visual feature is independent of other variables can be explained by models analogous to the colour triangle in colour discrimination. (shrink)

The appendages of the adult fruit fly and other insects and Arthropods develop from secondary embryonic fields that form after the primary anterior/posterior and dorsal/ventral axes of the embryo have been determined. In Drosophila, the position and fate of the different fields formed within each segment are determined by genes acting along both embryonic axes, within individual segments, and within specific fields. Since the major architectural differences between most Arthropod classes and orders involve variations in the number, type and (...) morphology of body appendages, the elucidation of the embryology and molecular genetics of the origin and patterning of insect limb fields may help to facilitate an understanding of both the mechanism of appendage formation and some of the major steps in the morphological evolution of the Arthropods. In this review, we will discuss recent studies that have advanced our understanding of both the origin and patterning of Drosophila leg and wing secondary fields. These results provide fresh insights into potentially general mechanisms of how body parts develop and evolve. (shrink)

In 1995, the eyeless (ey) gene was dubbed the “master‐regulator” of eye development in Drosophila. Not only is ey required for eye development, but its misexpression can convert many other tissues into eye, including legs, wings and antennae.(1) ey is remarkable for its ability to drive coordinate differentiation of the multiple cell types that have to differentiate in a very precise pattern to construct the fly eye, and for its power to override the previous differentiation programs of many other (...) diverse tissues. Even more remarkable, the ey homolog Pax6 and homologs of other eye determination genes from Drosophila are also required for eye development in vertebrates,(2,3) prompting reassessment of the evolution of vision throughout the animal kingdom.(4,5) Now Kumar and Moses have published a study that throws a new light on ey function in Drosophila.(6) According to their work, ey becomes a master regulator of eye development much later than previously thought, and is regulated by signalling through the Notch and EGFR signaling pathways. BioEssays 23:763–766, 2001. © 2001 John Wiley & Sons, Inc. (shrink)

“Looking for Asian America shows real people engaged in the full range of human activity. This is no small accomplishment for the photographer or his subjects. For Asian Americans it is extraordinary to be merely ordinary. To others, even if not to themselves, Asian Americans appear to be contradictions of identity—a Chinese-Yankee is a knockoff.” —Frank H. Wu, from the Foreword In search of contemporary Asian America, celebrated photographer Wing Young Huie—the only member of his family not born in (...) China—traveled with his wife Tara through nearly forty states to explore and document the funny, touching, and sometimes strange intersection of Asian American and American cultures. Looking for Asian America illustrates their rich and surprising journey across the United States. Through Huie’s eyes, keenly aware of his own Midwestern roots and perspective, we witness such images as a Vietnamese Elvis, Miss Congeniality on her cell phone in San Francisco’s Chinatown, a Hmong street sign in rural North Carolina, a meditating Falun Gong protestor in Washington, D.C., a bubble tea Valley Girl, and a Chinese theme park in Orlando. Huie’s camera captures ABCs, FOAs, and a self-described “redneck” Chinese restaurant owner near the Okefenokee Swamp. Taken together the photographs reveal a complex portrait of the U.S. cultural landscape, and their dignified elegance invites a closer, deeper look. Accompanied by the personal reflections of both Wing and Tara Huie, the nearly one hundred spectacular photos tell a story that both mirrors and contradicts stereotypes of Asian Americans, ultimately questioning what it means to be ethnic and American in the twenty-first century. Wing Young Huie has received widespread acclaim for his works, including Lake Street USA, documenting the cultural landscape of his native Minnesota. He is a recipient of a Bush Artist Fellowship and two-time recipient of the McKnight Photography Fellowship. He lives in Minneapolis. Frank H. Wu is dean of Wayne State University Law School and the author of Yellow: Race in America Beyond Black and White. Anita Gonzalez teaches in the Master of Liberal Studies Program at the University of Minnesota. (shrink)

In order to produce a coherent narrative, speakers must identify the characters in the tale so that listeners can figure out who is doing what to whom. This paper explores whether speakers use gesture, as well as speech, for this purpose. English speakers were shown vignettes of two stories and asked to retell the stories to an experimenter. Their speech and gestures were transcribed and coded for referent identification. A gesture was considered to identify a referent if it was produced (...) in the same location as the previous gesture for that referent. We found that speakers frequently used gesture location to identify referents. Interestingly, however, they used gesture most often to identify referents that were also uniquely specified in speech. Lexical specificity in referential expressions in speech thus appears to go hand‐in‐hand with specification in referential expressions in gesture. (shrink)

This Source Book is devoted to the purpose of providing such a basis for genuine understanding of Chinese thought (and thereby of Chinese life and culture, ...

The idea of a final end of human conduct – the highest good – lies at the centre of important parts of Kant’s philosophy, such as his moral theory, his philosophy of religion, his views on the historical progress of the human species, and his conception of human rationality. This collection of new essays attempts to re-evaluate the doctrine of the highest good and to determine its relevance for contemporary philosophy.

Sign languages modulate the production of signs in space and use this spatial modulation to refer back to entities—to maintain coreference. We ask here whether spatial modulation is so fundamental to language in the manual modality that it will be invented by individuals asked to create gestures on the spot. English speakers were asked to describe vignettes under 2 conditions: using gesture without speech, and using speech with spontaneous gestures. When using gesture alone, adults placed gestures for particular entities in (...) non-neutral locations and then used those locations to refer back to the entities. When using gesture plus speech, adults also produced gestures in non-neutral locations but used the locations coreferentially far less often. When gesture is forced to take on the full burden of communication, it exploits space for coreference. Coreference thus appears to be a resilient property of language, likely to emerge in communication systems no matter how simple. (shrink)

Anticipated nostalgia is a new construct that has received limited empirical attention. It concerns the anticipation of having nostalgic feelings for one’s present and future experiences. In three...

London Medical Group was founded in 1963. It was student-led, spawned Medical Groups in almost every UK medical school and met a need for non-partisan debate and dialogue in medical ethics. It became a victim of its own success as the Institute of Medical Ethics published the Pond Report in 1987, which recommended that medical ethics be incorporated into the undergraduate curriculum. Medical schools began to teach medical ethics and the General Medical Council demanded this in 1993's Tomorrow's Doctors. The (...) Institute of Medical Ethics had grown out of the LMG. After running a number of successful conferences for medical ethics teachers, the Institute of Medical Ethics is recapturing the natural innovative tendencies of students and junior doctors that the LMG and related Medical Groups had fostered. It is now launching itself as a membership organisation: the recommendations of the Francis report and responses to it have emphasised the need to support individuals with the ability, freedom and confidence to question the status quo from a reasoned ethical basis. The Institute of Medical Ethics aims to develop a robust medical ethics community ready to face the challenges of 21st Century healthcare. (shrink)