Obituary for Dr Mark Goodwin

The late Dr Mark Goodwin talking to Ethiopian farmers and children

The late Dr Mark Goodwin (1960-2018) talking to Ethiopian farmers and children

I am so saddened to write about the sudden death of my good friend, colleague and co-project leader, Mark Goodwin. We had very extensive common interests in the application and delivery of research for developing countries, research ethics, and in the role and importance of tertiary education and pedagogy. His inputs to my thinking about these areas have been critical to my research and how it impacts other people throughout the world, and I will so much miss the opportunities for discussions and working together in the future. His loss is an enormous shock: we spent much of his last day together discussing research and dissemination, and finalizing plans for his summit in Ethiopia the following week.

I first got to know Mark well 10 years ago in the context of the link with University of Gondar. He had real vision behind developing the PhD studentships to support staff members at the Ethiopian University. During previous visits to Ethiopia with Mark, we saw many scientific and cultural highlights together, all of which I am sure were applied in our research and teaching in the future (videos where we were together are on Youtube: Fishing tilapia in Lake Hawassa  or investigating conservation plantings to stop hillside erosion in Aksum). He brought extensive and insightful personal input to every one of us involved in the project, and was a true driver. I also saw a lot of Mark’s incredible help to individual students: in a time when so many want students to be considered as numbers and something to do form-filling correctness, he knew every single one he worked with, and would go out of his way to support them in any way he could, always working so the special talents of individuals came out. He challenged administration if needed or sorted out difficult issues. Examples include arranging for his project students to spend time with me in structured interviews, or finding ways to arrange suitable accommodation and subsistence payments, or working to obtain grants or funding to support visits.

Mark Goodwin (right) with Tesfeye Bizuayeu and Pat Heslop-Harrison in an Enset germplasm collection field

Mark Goodwin (right) with Tesfeye Bizuayeu and Pat Heslop-Harrison in an Enset germplasm collection field

In our current joint project on Abyssinian banana, enset or Ensete ventricosum, Mark, as co-Investigator, has been ensuring we go far beyond scientific goals towards building a group of collaborative stakeholders – including HEIs and established agricultural research organisations. The website is one of the outputs, with huge input to the contents and structure from Mark. Through Mark’s expertise, we could ensure our research aims were kept close to the agriculture, environmental, social and economic development issues, we kept engagement with the research, showed the necessary commitment to implementing the outcomes through collaborations, networking and negotiation.

I feel so fortunate to have spent a lot of time with Mark on his last day. We had lunch together in the University cafeteria, and had a typical wide-ranging discussions about delivery of impact for research projects and research assessment, the challenges he had just faced so successfully in leading the recruitment of Undergraduate students to the University, how we can ensure a high profile for the need for higher education for international development within governments and the UK, project supervision ideas and challenges, the latest news on our publications and so much more. We moved on to the topic of a meeting we had planned later in the afternoon: Mark was leading the organization of a summit and other meetings in Hawassa and Addis Ababa, Ethiopia, the following week. Mark was found in his office later that evening. He was looking forward to making inputs to the delivery of the development goals, and steering the scientific outputs to seeing how they could be used in education and dissemination, in policy and in application, in sustainable agriculture, economic or rural development, and environmental improvement.

Mark Goodwin with Worku Mhiret in the field in Ethiopia

Mark Goodwin with Worku Mhiret in the field in Ethiopia

I will certainly miss too discussions with Mark about wider issues of ethics, music and arts, global culture and history. He was truly a great friend, and one I could always rely upon. I know he will be missed by his many students and collaborators throughout the world.

Mark was taken from us far too early, but his contributions will live on through his huge impact to both research application and implementation, the development of higher education structures internationally, research evaluation and pedagogy, teaching in the UK and abroad, and the delivery of the projects for development.

Pat Heslop-Harrison

A notice about the sad loss of Mark is also published on the University of Leicester website, giving more details of his huge contributions to the Department of Genetics and Genome Biology and the School of Biological Sciences.




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Biodiversity in Ethiopian linseed: molecular characterization of landraces

342. Mhiret WN, Heslop-Harrison JS. 2018. Biodiversity in Ethiopian linseed (Linum usitatissimum L.): molecular characterization of landraces and some wild species. Genetic Resources and Crop Evolution 65: 1603–1614. or author version: Linseed Linum Ethiopia Molecular Diversity Worku Mhiret GRACE 2018 Author Version

Linseed molecular biodiversity and variation. Negash and Heslop-Harrison 2018 GRACE Genet Res Crop Evol 65: 1603

Molecular characterization of germplasm is important for sustainable exploitation of crops. DNA diversity was measured using inter-retrotransposon-amplified-polymorphism and inter-simple-sequence-repeat markers in 203 Ethiopian landraces and reference varieties of linseed (flax, Linum usitatissimum) and wild Linum species. Molecular diversity was high (PIC, 0.16; GD, 0.19) compared to other reports from the species. Genotyping separated reference from landrace accessions, and clustered landrace accessions from different altitudes and geographical regions. Collections showed evidence for recent introduction of varieties in some regions. The phylogeny supported L. bienne Mill. as the progenitor of domesticated L. usitatissimum. Markers developed here will be useful for genetic mapping and selection of breeding lines. The results show the range of characters that can be exploited in breeding lines appropriate for smallholder and commercial farmers in Ethiopia, producing a sustainable, secure, high-value crop meeting agricultural, economic and cultural needs.

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ImmunoFISH – Simultaneous visualisation of proteins and DNA sequences gives insight into meiotic processes in nuclei of grasses

Sepsi A, Fábián A, Jäger K, Heslop-Harrison JS, Schwarzacher T. ImmunoFISH: simultaneous visualisation of proteins and DNA sequences gives insight into meiotic processes in nuclei of grasses. Frontiers in Plant Science. 9: 1193.

Combined Immunolabelling and In Situ Hybridization FISH. Sepsi et al. 2018. Frontiers Plant Sci 9: article 1193

Combined immunolabelling and in situ hybridization FISH. Sepsi et al. 2018. Frontiers Plant Sci 9: article 1193

ImmunoFISH is a method combining immunolabelling (IL) with fluorescent in situ hybridisation (FISH) to simultaneously detect the nuclear distribution of proteins and specific DNA sequences within chromosomes. This approach is particularly important when analysing meiotic cell division where morphogenesis of individual proteins follows stage-specific changes and is accompanied by a noticeable chromatin dynamism. The method presented here is simple and provides reliable results of high quality signal, low background staining and can be completed within 2 days following preparation. Conventional widefield epifluorescent or laser scanning microscopy can be used for high resolution and three-dimensional analysis. Fixation and preparation techniques were optimised to best preserve nuclear morphology and protein epitopes without the need for any antigen retrieval. Preparation of plant material involved short cross-linking fixation of meiotic tissues with paraformaldehyde (PFA) followed by enzyme digestion and slide-mounting. In order to avoid rapid sample degradation typical of shortly fixed plant materials, and to be able to perform IL later, slides were snap-frozen and stored at -80C. Ultra-freezing produced a remarkable degree of structural preservation for up to 12 months, whereby sample quality was similar to that of fresh material. Harsh chemicals and sample dehydration were avoided throughout the procedure and permeability was ensured by a 0.1–0.3% detergent treatment. The ImmunoFISH method was developed specifically for studying meiosis in Triticeae, but should also be applicable to other grass and plant species.


Sepsi A, Fábián A, Jäger K, Heslop-Harrison JS, Schwarzacher T. ImmunoFISH: simultaneous visualisation of proteins and DNA sequences gives insight into meiotic processes in nuclei of grasses. Frontiers in Plant Science. 9: 1193.



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An Immortalized Genetic Mapping Population for Perennial Ryegrass: A Resource for Phenotyping and Complex Trait Mapping

Velmurugan J, Milbourne D, Connolly V, Heslop-Harrison JS, Anhalt UC, Lynch MB, Barth S. 2018. An immortalized genetic mapping population for perennial ryegrass: a resource for phenotyping and complex trait mapping. Frontiers in Plant Science 9: article 717

Lolium Genetic Map and RI lines showing population structure. Velmurugan et al. 2018. Frontiers in Plant Science 9: article 717

Lolium Genetic Map and RI lines showing population structure. Velmurugan et al. 2018. Frontiers in Plant Science 9: article 717

To address the lack of a truly portable, universal reference mapping population
for perennial ryegrass, we have been developing a recombinant inbred line (RIL)
mapping population of perennial ryegrass derived via single seed descent from a
well-characterized F2 mapping population based on genetically distinct inbred parents
in which the natural self-incompatibility (SI) system of perennial ryegrass has been
overcome. We examined whether it is possible to create a genotyping by sequencing
(GBS) based genetic linkage map in a small population of the F6 generation of this
population. We used 41 F6 genotypes for GBS with PstI/MspI-based libraries. We
successfully developed a genetic linkage map comprising 6074 SNP markers, placing a
further 22080 presence and absence variation (PAV) markers on the map. We examined
the resulting genetic map for general and RIL specific features. Overall segregation
distortion levels were similar to those experienced in the F2 generation, but segregation
distortion was reduced on linkage group 6 and increased on linkage group 7. Residual
heterozygosity in the F6 generation was observed at a level of 5.4%. There was a high
proportion of chromosomes (30%) exhibiting the intact haplotype of the original inbred
parents of the F1 genotype from which the population is derived, pointing to a tendency
for chromosomes to assort without recombining. This could affect the applicability
of these lines and might make them more suitable for situations where repressed
recombination is an advantage. Inter- and intra-chromosomal linkage disequilibrium
(LD) analysis suggested that the map order was robust. We conclude that this RIL
population, and subsequent F7 and F8 generations will be useful for genetic analysis
and phenotyping of agronomic and biological important traits in perennial ryegrass.

Click to access fpls-09-00717.pdf

Keywords: perennial ryegrass, Lolium perenne, recombinant inbred lines (RIL), genotyping by sequencing, mapping population, phenotyping

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Herbert Macgregor 1933-2018: a personal tribute

Topsham Church and Professor Herbert Macgregor's Memorial Service

Topsham Church and the Service of Thanskgiving for the Life of Professor Herbert Macgregor

My Tribute to Professor Herbert Macgregor (22nd April 1933 – 22nd July 2018) delivered at his Thanksgiving Service on 13th August 2018

I am humbled to be here today to pay tribute to the wonderful scientist, leader and mentor, Herbert Macgregor. As a fellow cytogeneticist, Herbert has been right at the top of my field, and despite the great sadness, I am happy to be able to give you some thoughts about his remarkable and productive life. I first came across Herbert’s name in my earliest undergraduate studies with other cytogeneticists, Carl Swanson and Hugh Rees, learning of Herbert’s discovery (with Mick Callan) of the fact that each of the chromosomes within a cell nucleus – 46 of them in the case of human – contained a single DNA molecule, 100s of millions of DNA bases long, from end to end (Callan and Macgregor, 1958). Today, this is such a tenet of cell biology and genomics that it is accepted as ‘obvious’: the importance of the result supported by detailed experimental work stands equal with the other key discoveries of the late 1950s and 1960s, including the DNA helix, the genetic code, transcription, and semi-conservative replication of DNA. Herbert was a pioneer with techniques, using them and developing model systems to address many novel questions throughout his career. His book from the early 1980s, “Working with Animal Chromosomes” is still the standard reference for chromosome preparations and regularly used in my lab, while his 1993 volume, Introduction to Cytogenetics, is another standard science for interpreting chromosome form and behaviour, and appreciating its evolutionary significance.

I first met Herbert at a chromosome conference in 1982 organized in Lübeck, Germany. Herbert’s talk was a highspot of the meeting – one of those annotated with three stars in my programme. Like so many of Herbert’s papers, the title is as thoughtful and current now as when the paper was presented and written up, “The evolutionary consequences of major genomic changes”: remember this was long before the science of genomics was the huge industry that it has become. Other papers had titles such as “High resolution map” or “Application of the technique”, or discussed karyotypes and chromosome morphology. Herbert’s paper, based on his own detailed investigations of the chromosomes in one of his favoured groups, the newts and other amphibians, makes conclusions and generates ideas that are still current and important today.

Two years later, Herbert and Alma organized an important meeting of the British Society of Developmental Biology, BSDB, in Leicester, where Herbert was Professor and Head of the Department of Zoology from 1970. This was one of my first conference talks, and I remember well the exceptional organization and range of exciting talks on the topic “Genes, chromosomes and computer models in developmental biology”. At that time 34  years ago, “Computer models”  were unheard of in chromosome biology, but the contents of the volume are truly visionary. Herbert and Alma’s words in the introduction, “the frontier of this symposium lies between the biologists and computer scientists, between biology and biochemistry and mathematics and modelling” well summarize the range of Herbert’s research and its implications for the rest of his career. This 1984 meeting was my first visit to Leicester where I am now based, and there were some strange events around the conference. I remember ending up in a pub with a number of the most notable conference participants, including Adrian Bird and Chris Bostock, and witnessing still the most vicious fight I have ever seen, as well as one or two unusual conference participants!

I continued to follow with much excitement Herbert’s work, now linking gene expression with its physical manifestation as loops coming out from a special type of chromosome – lampbrush chromosomes – in amphibians and birds. This research on the arrangement and expression of DNA sequences in the genomes had major implications for molecular biology and the functioning of cell nuclei, as well as for evolution and developmental biology. Once again, this research is now part of the unquestioned canon of scientific knowledge. All through his work, from his very first papers in the late 1950s, Herbert was able to integrate microscopy with molecular biology – something that is today even more important than it was 50 years ago.

I got to know Herbert much better from the early 1990s, when he invited me to join him in a new publishing venture, setting up the Journal Chromosome Research, now in its 25th successful year. With Herbert’s vision (set out in his Macgregor1993_Article_Editorial, followed up by the insightful commentary, from 1993, on Macgregor1993_Article_ChromosomeResearchLookForwardT), it has gained a distinctive niche in a publishing area, long before the huge expansion in numbers of publications the last decade. I gained enormously from seeing his firm but helpful interactions with the series of publishers we worked through, and carefully thought-through handling of the interactions with authors and editors to build the reputation of the journal. We were now in close contact, not only with publishing but with respect to our research and the chromosome research environment. So I discussed with him about 1999 that my then institute was in an uncomfortable state of upheaval, and about any academic positions coming up? Retiring around that time, but still maintaining his lab., he asked the right people and soon after, a rather specialized advertisement appeared in a newspaper, sandwiched between adverts for the chief fire officer for B&Q (a chain of UK DIY shops), and head groundsman for the Leicester City Football Club. So I moved to Leicester, and his wise advice was so valuable to me. Most notably, Herbert valued the exceptional people he worked with, particularly among the support staff. He invariably appreciated their immense contributions, showing the importance of their roles, and the impact they could have. In this way, he was able to build an extraordinary and productive world-class Department, and I saw the remaining atmosphere he fostered in Leicester. Within the environment he fostered in his Department, he had no truck with nonsense from administrators and senior colleagues, and was forthright in his robust views. I should also add that he held the most wonderful Christmas parties, with several dozen people from the wider Department.

Herbert cared deeply about his teaching and undergraduates and gave me extremely valuable advice as I took over in teaching some of the introductory genetics and biology courses. His lecturing style was immensely approachable but still challenging, and I was not surprised to hear he continued outreach and learning activities through U3A here in Exeter. Among other things, he maintained a remarkable website here at the University of Exeter on lampbrush chromosomes, providing a resource for all – not least with its progressive CC-BY open accessibility without copyright.

Personally, it was always wonderful and memorable to meet Herbert. At the time of moving to Leicester, we stayed with him and Alma, at his farm, The Leys. He had many musical, technical and nautical interests that were always a pleasurable digression to discuss. So many characteristics of Herbert come through from every interaction I had with him, and I am very happy to have been able to pay this tribute. Hebert was, or course, very proud of his Scottish Heritage, and retained his musical accent that was so easy to listen to, carrying erudition and distinction in its tones. I do remember a meeting that ended with a Burns Night supper, where we had the privilege of hearing Herbert giving the most authentic recitation of the words of The Bard.

So I will end with some other words that are appropriate from Robbie Burns’ “Epitaph on my Own Friend”:

An honest man here lies at rest,
As e’er God with His image blest:
The friend of man, the friend of truth;
The friend of age, and guide of youth:
Few hearts like his, with virtue warm’d,
Few heads with knowledge so inform’d:
If there’s another world, he lives in bliss;
If there is none, he made the best of this.

Pat Heslop-Harrison.

Mine was one of four Tributes delivered at Professor Macgregor’s Thanksgiving Service. Others covered his early years including National Service, his Family, and life during his retirement in Topsham, Exeter.

Thanks to Dr Alma Swan, Herbert’s publication list is posted Microsoft word verstion HerbertMacgregor_PublicationList_PUBLISHED WORK  and given below:


 1958 to 2012

  1. Callan, H.G. and H.C. Macgregor (1958). Action of deoxyribonuclease on lampbrush chromosomes. Nature, 181: 1479-1480.
  2. Macgregor, H.C. (1962). The behaviour of isolated nuclei. Exp.Cell Res., 26: 520-525.
  3. Macgregor, H.C. and H.G. Callan (1962). The actions of enzymes on lampbrush chromosomes. Quart. J. Microsc. Sci.,  103:  172-203.
  4. Macgregor, H.C. (1963) Morphological variability and its physiological origin in oocyte nuclei of the crested newt. Quart.J. Microsc. Sci., 104:  351-368.
  5. Macgregor, H.C. and T.M. Uzzell (1964) Gynogenesis in salamanders related to Ambystoma jeffersonianum. Science, 143: 1043-1045.
  6. Macgregor, H.C. and P.A. Thomasson(1965). The fine structure of two archigregarines, Selenidium fallax and Ditrypanocystis cirratuli.  J.Protozool. 12:  438-443.
  7. Macgregor, H.C. (1965). The role of lampbrush chromosomes in the formation of nucleoli in amphibian oocytes. Quart. J. Microsc. Sci., 106: 215-228.
  8. Macgregor, H.C. (1965). The effects of mammalian gonadotrophins on the yolky oocytes of crested newts. Arch.Anat.Micr. 54:  652-653.
  9. Macgregor, H.C. (1967). Pattern of incorporation of 3H uridine into RNA of amphibian oocyte nucleoli. J.Cell.Sci. 2: 145-150.
  10. Macgregor, H.C. and J.B. Mackie (1967). The fine structure of the cytoplasm in salivary glands of Simulium. J. Cell. Sci. 2: 137-144.
  11. Perkowska, E., H.C. Macgregor and  M.L. Birnstiel (1968). Gene amplification in the oocyte nucleus of mutant and wild type Xenopus laevis. Nature, 217: 649-650.
  12. Walker, M. and H.C. Macgregor (1968). Spermatogenesis and the structure of the mature sperm in Nucella lapillus (L). J.Cell.Sci. 3: 95-104.
  13. Macgregor, H.C., (1968). Nucleolar DNA in oocytes of Xenopus laevis. J. Cell Sci.,  3:  437-444.
  14. Gall, J.G., H.C. Macgregor, and M.E. Kidston(1969). Gene amplification in the oocytes of dytiscid water beetles. Chromosoma, 26:169-187.
  15. Riemann, W., C. Muir, and H.C. Macgregor (1969). Sodium and potassium in oocytes of Triturus cristatus. J.Cell Sci.,  4: p. 299-304.
  16. Macgregor, H.C. (1969). Observations on the role of the germinal vesicle in amphibian and insect oogenesis. Heredity, 24: p. abstract in Proceedings of the 159th meeting of the Genetical Soc. of G.B.
  17. Macgregor, H.C. and J. Kezer (1970). Gene amplification in oocytes with 8 germinal vesicles from the tailed frog Ascaphus truei Stejneger. Chromosoma, 29: 189-206.
  18. Macgregor, H.C. and H. Stebbings (1970). A massive system of microtubules associated with cytoplasmic movement in telotrophic ovarioles. J.Cell Sci., 6: 431-449.
  19. Macgregor, H.C. and S.J. Moon (1971). Some measurements on amphibian oocyte nucleoli. Z.  Zellforsch., 122:  273-282.
  20. Kezer, J. and H.C. Macgregor (1971). A fresh look at meiosis and centromeric heterochromatin in the red-backed salamander Plethodon c. cinereus (Green). Chromosoma, 33:  146-166.
  21. Macgregor, H.C. and J. Kezer (1971). The chromosomal localisation of a heavy satellite DNA in the testis of Plethodon c. cinereus . Chromosoma, 33: 167-182.
  22. Kezer, J., H.C. Macgregor, and E. Schabtach (1971). Observations on the membranous components of amphibian oocyte nucleoli. J. Cell Sci. 8: 1-17.
  23. Macgregor, H.C. and M. Vlad (1972). Interlocking and knotting of ring nucleoli in amphibian oocytes. Chromosoma, 39: 205-214.
  24. Macgregor, H.C. (1972). The nucleolus and its genes in amphibian oogenesis. Biol.Rev. 47: 177-210.
  25. Macgregor, H.C. (1973). Amplification, polytenisation and nucleolus organisers. Nature New Biology, 246: 81-82.
  26. Macgregor, H.C., H.A. Horner, C.A. Owen and I. Parker (1973). Observations on centromeric heterochromatin and satellite DNA in salamanders of the genus Plethodon. Chromosoma, 43: 329-348.
  27. Macgregor, H.C. and M. Walker (1973). The arrangement of chromosomes in nuclei of sperm from plethodontid salamanders. Chromosoma, 40: 243-262.
  28. Macgregor, H.C. and J. Kezer (1973). The nucleolar organiser of Plethodon c. cinereus. I. Location of the organiser by in-situ nucleic acid hybridisation. Chromosoma, 42:  415-426.
  29. Kezer, J. and H.C. Macgregor (1973). The nucleolar organiser of Plethodon c. cinereus. II. The lampbrush nucleolar organizer. Chromosoma, 42: 427-444.
  30. Walker, M.H. and H.C. Macgregor (1974). The arrangement of chromosomes in elongate sperm heads. in  Chromosomes Today, 5 :  13 – 22.  (John Wiley & Sons).
  31. Mizuno, S. and H.C. Macgregor (1974). Chromosomes, and sequences and evolution in salamanders of the genus Plethodon. Chromosoma, 48: p. 239-296.
  32. Macgregor, H.C., S. Mizuno, and M. Vlad. Some recent studies on chromosomes and DNA sequences in salamanders. in Chromosomes Today 5 : 331 – 339. (John Wiley & Sons).
  33. Vlad, M. and H.C. Macgregor (1975). Chromomere number and its genetic significance in lampbrush chromosomes. Chromosoma, 50: 327-347.
  34. Hennen, S., S. Mizuno, and H.C. Macgregor (1975). In-situ hybridisation of rDNA labelled with 125-I to metaphase and lampbrush chromosomes of newts. Chromosoma, 50:340-360.
  35. Macgregor, H.C., M. Vlad, and L. Barnett (1976). An investigation of some problems concerning nucleolus organisers in salamanders. Chromosoma, 59: 283-299.
  36. Macgregor, H.C. and S. Mizuno (1976). In-situ hybridisation of ‘nick-translated’ 3H-ribosomal DNA to chromosomes from salamanders. Chromosoma, 54:  15-25.
  37. Mizuno, S., C. Andrews, and H.C. Macgregor (1976) Interspecific ‘common’ DNA sequences in salamanders of the genus Plethodon. Chromosoma, 58: 1-31.
  38. Macgregor, H.C. and C. Andrews (1977). The arrangement and transcription of ‘middle repetitive’ DNA sequences on lampbrush chromosomes of Triturus. Chromosoma, 63: 109-126.
  39. Macgregor, H.C. (1977). Lampbrush Chromosomes. in Chromatin and Chromosome Structure., R.A. Eckhardt and Hseuh-Jei Li (Editors). pp. 339 – 357.  Academic Press:
  40. Macgregor, H.C. and C. Jones (1977). Chromosomes, DNA sequences, and evolution in salamanders of the genus Aneides. Chromosoma, 63: 1-9.
  41. Macgregor, H.C. and G.T. Morgan (1978). Repetitive DNA and chiasma failure in chromosome I of crested newts. J.Cell Biol., 79: 133a. (Abstract)
  42. Macgregor, H.C. (1978). Trends in the evolution of very large chromosomes. Proc.Roy.Soc.London, B 283: 309-318.
  43. Morgan, G.T., H.C. Macgregor, and A. Colman (1979). Multiple ribosomal sites revealed by in situ hybridisation of Xenopus  rDNA to Triturus  lampbrush chromosomes. Chromosoma,   80: 309-330.
  44. Macgregor, H.C. and S. Sherwood (1979). The nucleolus organisers of Plethodon and Aneides  located by in-situ  nucleic acid hybridisation with Xenopus  3H ribosomal RNA. Chromosoma, 72:  271-280.
  45. Macgregor, H.C. (1979). In-situ hybridisation of highly repetitive DNA to chromosomes of Triturus cristatus. Chromosoma, 71:  57-64.
  46. Macgregor, H.C. and L. Klosterman (1979). Observations on the cytology of Bipes (Amphisbaenia) with special reference to its lampbrush chromosomes. Chromosoma, 72: 67-87.
  47. Macgregor, H.C. (1980). Recent developments in the study of lampbrush  chromosomes. Heredity, 44: 3-35.
  48. Varley, J.M., H.C. Macgregor, and H.P. Erba (1980). Satellite DNA is transcribed on lampbrush chromosomes. Nature, 283:  686-688.
  49. Hill, R.S. and Macgregor, H.C. (1980). The development of lampbrush chromosome-type transcription in the early diplotene oocytes of Xenopus laevis: an electron microscope analysis.  J. Cell Sci.  44, 87 – 101.
  50. Macgregor, H.C., J.M. Varley, and G.T. Morgan (1981). The transcription of satellite and ribosomal DNA sequences on lampbrush chromosomes of crested newts., in International Cell Biology. Springer-Verlag: p. 33-46.
  51. Macgregor, H.C. and H.A. Horner (1980). Heteromorphism for chromosome I, a requirement for normal development in crested newts. Chromosoma, 76: 111-122.
  52. Macgregor, H.C. (1980). Recent developments in the study of lampbrush chromosomes. Heredity, 44:  3-35.
  53. Varley, J. M., H.C. Macgregor,  I. Nardi,  C. Andrews and H.P. Erba (1980). Transcription of highly repeated DNA sequences during the lampbrush stage in Triturus cristatus carnifex. Chromosoma, 80: 289-307.
  54. Macgregor, H.C. (1982). Big chromosomes and speciation amongst Amphibia.  in Genome Evolution, G.A. Dover and R.B. Flavell (Editors). Academic Press: pp. 325-341.
  55. Macgregor, H.C. (1982). Ways of amplifying ribosomal genes.  in  The Nucleolus.  Society for Experimental Biology Seminar Series 15, E.G. Jordan and C.A. Cullis (Editors). Cambridge University Press: Cambridge. pp. 129-152.
  56. Macgregor, H.C. and E. del Pino (1982). Ribosomal gene amplification in  ultinucleate oocytes of the egg brooding hylid frog Flectonotus pygmaeus. Chromosoma, 85:  475-488.
  57. Horner, H.A. and H.C. Macgregor (1983). C value and cell volume; their significance in the evolution and development of amphibians. J.Cell Sci., 63: 135-146.
  58. Macgregor, H.C. and J.M. Varley (1983). Working with Animal Chromosomes. John Wiley and Sons Ltd.: Chichester and New York.
  59. Macgregor, H.C., Horner, H.A. and Sims, S.H. (1983). Newt chromosomes and some problems in evolutionary cytogenetics.  Proceedings of the Kew Chromosome Conference II, Allen and Unwin.  pp. 283 – 294.
  60. Swan, A.P., H.C. Macgregor, and R. Ransom (editors) (1984). Programmes for Development.  Symposium of the British Society for Developmental Biology. 1984. The Company of Biologists.
  61. Macgregor, H.C. (1984). The lampbrush chromosomes of animal oocytes.  in Chromosome Structure and Function, M. Risley, (Editor). van Rostrand and Reinhold Publishing Corp: New York. pp. 152 – 186.
  62. Macgregor, H.C. and A.P. Swan (1984). An introduction to Programmes for Development. J.Embryol.Exp.Morph. 83(Supplement):  1-6.
  63. Sims, S.H., H.C. Macgregor, P.A. Pellatt and H.A. Horner (1984). Chromosome 1 in crested and marbled newts genus Triturus: an extraordinary case of  heteromorphism and independent chromosome evolution. Chromosoma, 89: 169-185.
  64. Macgregor, H.C. (1984). Lampbrush chromosomes and gene utilisation in meiotic prophase. In Controlling Events in Meiosis, W. Evans and H.G.Dickinson (Editors).  The Company of Biologists. pp. 333 – 348.
  65. Macgregor, H.C. (1984). The evolutionary consequence of major genomic changes in Amphibia. In Chromosomes Today, 8 M.D.Bennett, A. Gropp, U. Wolf (Editors).  George Allen and Unwin.  pp. 256 – 267.
  66. Horner, H.A. and H.C. Macgregor (1985). Normal development in newts (Triturus) and its arrest as a consequence of an unusual chromosomal situation.  J. Herpetology, 19:  216 – 270.
  67. Baldwin, L. and H.C. Macgregor (1985). Centromeric satellite DNA in the newt Triturus cristatus karelinii and related species: its distribution and transcription on lampbrush chromosomes. Chromosoma, 92: 100-107.
  68. Macgregor, H.C. and S.K. Sessions (1986). The biological significance of variation in satellite DNA and heterochromatin in newts of the genus Triturus: an evolutionary perspective., in The Evolution of DNA Sequences, B.C. Clarke, A. Robertson and A.J. Jeffreys (Editors). Phil. Trans. R. Soc. Lond.: B 312:  53 – 70.
  69. Macgregor, H.C. (1986). Care, maintenance and captive breeding of newts (Triturus).  In UFAW Handbook, Longman.  pp. 768 – 772.
  70. Schmid, M., S.H. Sims, T. Haaf and H.C. Macgregor (1986). Chromosome banding in Amphibia X. 18S and 28S ribosomal RNA genes, nucleolus organizers and nucleoli in Gastrotheca riobambae .   Chromosoma, 94:  139-145.
  71. Macgregor, H.C. and S.K. Sessions (1986). Models for evolution of large genomes and karyotypes of urodeles. Verhandlungen der Duetzchen Zoologischen Gesellschaft., 79: 137-148.
  72. Macgregor, H.C. (1986). The lampbrush chromosomes of animal oocytes. in Chromosome Structure and Function, M.Risley (Editor), Van Rostrand & Reinhold Publishing Corporation, New York.: pp. 152-186.
  73. Macgregor, H.C. (1987). Lampbrush Chromosomes. J. Cell. Sci. 88: 7-9.
  74. Macgregor, H.C. (1988). The Evolutionary Cytogenetics of Triturus (Amphibia, Urodela):  An overview. In Symposium on the Evolution of Terrestrial Vertebrates, G. Chiara (Editor).  Mucchi, Modena.  pp. 153 – 170.
  75. Macgregor, H.C. and J. Varley (1988). Working with Animal Chromosomes. 2nd edition. John Wiley & Sons.
  76. Macgregor, H.C. (1988). Chromosome heteromorphism in newts (Triturus) and its significance in in relation to evolution and development. In Amphibian Cytogenetics and Evolution, D. Green and S.K.Sessions (Editors). Academic Press.  pp. 175 – 196.
  77. Sessions, S.K., H.C. Macgregor, M. Schmid and T. Haaf (1988). Cytology, embryology and evolution of the developmental arrest syndrome in newts of the genus Triturus (Caudata: Salamandridae).  J. exp. Zool.  248:  321 – 334.
  78. Macgregor, H.C. (1990). Newts and two studies in molecular cytogenetics., in Cytogenetics of Amphibians and Reptiles, E. Olmo (Editor).  Birkhauser Verlag. pp. 61-84.
  79. Varley, J.M., H.C. Macgregor and L. Barnett (1990). Characterization of a short, highly repeated and centromerically localized DNA sequence in crested and marbled newts of the genus Triturus.  Chromosoma, 100:  15 – 31.
  80. Macgregor, H.C., S.K. Sessions, and J.W. Arntzen (1990). An integrative analysis of phylogenetic relationships amongst newts of the genus Triturus (family Salamandridae) using comparative biochemistry, cytogenetics and reproductive interactions. J.Evol.Biol. 3:  329-374.
  81. Solovei, I., E. Gaginskaya, T.D. Allen and H.C. Macgregor (1992). A novel structure associated with a lampbrush chromosome in the chicken, Gallus domesticus. J. Cell Science, 101: 759-772.
  82. Macgregor, H.C. (1992). Introduction to Biological Sciences. in The Pergamon Encyclopedia for Higher Education. 4:  2181 – 2183.  Pergamon Press, Oxford.
  83. Macgregor, H.C. (1992). Evolutionary Biology. In Pergamon Encyclopedia for Higher Education, 4: 2226 – 2231.  Pergamon Press, Oxford.
  84. Solovei, I., E. Gaginskaya, N. Hutchison and H.C. Macgregor  (1993).  Avian sex chromosomes in the lampbrush form:  the ZW lampbrush bivalents from six species of bird. Chromosome Research, 1:  153-166.
  85. Macgregor, H.C. (1993). British Universities in the World of Business. The New Academic, 3:  12-14.
  86. Macgregor, H.C. (1993). An Introduction to Animal Cytogenetics. Chapman and Hall: London.
  87. Macgregor, H.C. (1993). Chromosome Research – look forward to 2001.  Chromosome Research, 1: 5-7
  88. Solovei, I., E.R. Gaginskaya, and H.C. Macgregor (1994). The arrangement and transcription of telomere DNA sequences at the ends of lampbrush chromosomes of birds. Chromosome Research, 2: 460-470.
  89. Fairchild, P.J., Macgregor H.C. (1994). Asymmetric loops. Current Biology, 4: 919.
  90. Solovei, I., H.C. Macgregor and E. Gaginskaya (1995). Single stranded nucleic acid binding structures on chicken lampbrush chromosomes.  J. Cell Sci.  108, 1391 – 1396.
  91. Solovei, I., H.C. Macgregor and E. Gaginskaya (1995). Specifically terminal clusters of telomere DNA sequences are transcribed from the C-rich strand on chicken lampbrush chromosomes.  Proc. Kew Chromosome Conference IV.  P.F.Brandham and M.D.Bennett (eds. Royal Botanic Gardens, Kew, Press. 1995.  pp.  323 – 330.
  92. Macgregor, H.C. (1995). Crested Newts: Ancient Survivors.  British Wildlife 7:  1 – 8.
  93. Macgregor, H.C. (2011). From Bones to Biotechnology: 50 years of new biology in the Old World.   In A History of the University in Europe (ed. Walter Ruegg) Volume IV, 451 – 471. Cambridge University Press.
  94. Hori, T., Susuki, Y., Solovei, I., Saitoh, Y., Hutchison, N., Ikeda, Joh-E., Macgregor, H.C., Mizuno, S. (1996). Characterization of DNA sequences constituting the terminal heterochromatin of the chicken Z chromosome. Chromosome Research  4 :  411 – 426.
  95. Joffe, B.I., Solovei, I. V., Macgregor, H.C. (1996) Ends of chromosomes in Polycelis tenuis (Platyhelminthes) have telomere repeat TTAGGG. Chromosome Research  4:  323-324.
  96. Macgregor H.C., Solovei I., and Mizuno, S. (1997)  Lampbrush chromosomes as systems for high resolution studies of meiotic chromosome structure.  in  Chromosome Segregation and Aneuploidy, (ed. A. Abbondandolo, B.K.Vig and R. Roi). European Commission Joint Research Centre Publication. pp 172 – 183.
  97. Solovei, I., Joffe, B. Gaginskaya, E., Macgregor, H.C. (1996) Transcription on lampbrush chromosomes of a centromerically localized highly repeated DNA in pigeon (Columba) relates to sequence arrangement. Chromosome Research 4, 588 – 603.
  98. Ogawa, A., Solovei, I., Hutchison, N., Saitoh, Y., Ikeda, J-E., Macgregor, H.C., Mizuno, S. (1997).  Molecular characterization and cytological mapping of a non-repetitive DNA sequence region from the W chromosome of chicken and its use as a universal probe for sexing Carinatae birds.  Chromosome Research  5,  93 – 101.
  99. Joffe, B., Solovei, I., Macgregor,  H.C. (1998)  Ordered arrangement and rearrangement of chromosomes during spermatogenesis in two species of planarians (Platyhelminthes).  Chromosoma 107, 173 – 183.
  100. Solovei, I., Joffe, B., Hori, T., Thomson, P., Mizuno, S., Macgregor, H.C. (1998) Unordered arranmgement of chromosomes in nuclei of chicken spermatozoa. Chromosoma  107,  184 – 188.
  101. Mizuno, S., Macgregor H.C. (1998) The ZW lampbrush chromosomes of birds: a unique opportunity to look at the molecular cytogenetics of sex chromosomes.  Cytogenet. Cell Genet. 80, 149 – 157.
  102. Solovei, I., Ogawa, A., Naito, M., Mizuno, S., Macgregor H.C. (1998) Specific chromomeres on the chicken W lampbrush chromosome contain specific repetitive DNA sequence families. Chromosome Research  6, 323 – 327.
  103. Baker, S., Greig, C., Macgregor, H.C., Swan, A. (1998) Exmoor ponies:  Britain’s prehistoric wild horses?  British Wildlife  9, 304 – 313.
  104. Macgregor, H.C. (2000) The Future of Chromosomes. In Chromosomes Today, (E. Olmo and C.A.Redi, eds.) 13, 305 – 313.  Birkhauser Verlag.
  105. Teranishi, M., Shimada, Y., Hori, T., Nakabayashi, O., Kikuchi, T., MacLeod, T., Pym, R., Sheldon, B., Solovei, I., Macgregor, H.C., Mizuno, S. (2001) Transcripts of the MHM region on the chicken Z chromosome accumulate as non-coding RNA in the nucleus of female cells adjacent to the DMRT1 locus. Chromosome Research, 9: 147 – 165.
  106. Sessions, S.K., Macgregor, H.C. (2009) The necessity of Darwin. Chromosome Research 17: 437 – 442.
  107. Macgregor, H.C. (2012) Chromomeres revisited. Chromosome Research 20: 911 – 924.
  108. Macgregor, H.C. (2012) So what’s so special about these things called lampbrush chromosomes? Chromosome Research 20: 903 – 904.



Edits 14/8/18: hyperlinks, explain B&Q as shop.

Edit 10/9/18: Publication List for Professor HC Herbert Macgregor added


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Complete mitogenomes from Kurdistani sheep – abundant centromeric nuclear copies representing diverse ancestors

Kurdistani sheep breeds and part of their mitochondrial genome From Mustafa et al. Mitochondrial DNA part A 2018.

Kurdistani sheep breeds and part of their mitochondrial genome From Mustafa et al. Mitochondrial DNA part A 2018.

341. Mustafa SI, Schwarzacher T, and Heslop-Harrison JS. 2018. Complete mitogenomes from Kurdistani sheep: abundant centromeric nuclear copies representing diverse ancestorsMitochondrial DNA Part A (publisher – see free publisher link below if you can’t access) AND Mustafa et al_2018 AuthorVersion

The geographical center of domestication and species diversity for sheep (Ovis aries) lies around the Kurdistan region of Northern Iraq, within the ‘Fertile Crescent’. From whole genome sequence reads, we assembled the mitochondrial genomes (mtDNA or mitogenome) of five animals of the two main Kurdistani sheep breeds Hamdani and Karadi and found they fitted into known sheep haplogroups (or matrilineages), with some SNPs. Haplotyping 31 animals showed presence of the main Asian (hpgA) and European (hpgB) haplogroups, as well as the rarer Anatolian haplogroup hpgC. From the sequence reads, near-complete genomes of mitochondria from wild sheep species (or subspecies), and even many sequences similar to goat (Capra) mitochondria, could be extracted. Analysis suggested that these polymorphic reads were nuclear mitochondrial DNA segments (numts). In situ hybridization with seven regions of mitochondria chosen from across the whole genome showed strong hybridization to the centromeric regions of all autosomal sheep chromosomes, but not the Y. Centromeres of the three submetacentric pairs and the X chromosomes showed fewer copies of numts, with varying abundance of different mitochondrial regions. Some mitochondrial-nuclear transfer presumably occurred before species divergence within the genus, and there has been further introgression of sheep mitochondrial sequences more recently. This high abundance of nuclear mitochondrial sequences is not reflected in the whole nuclear genome assemblies, and the accumulation near major satellite sequences at centromeres was unexpected. Mitochondrial variants including SNPs, numts and heteroplasmy must be rigorously validated to interpret correctly mitochondrial phylogenies and SNPs.

Keywords: nuclear mitochondrial DNA segments (numts); mitogenome diversity; massively parallel sequencing; fluorescent in situ hybridization; sheep domestication

Free link to paper from publishers: (limited to 50 uses so please look at author version first or use your academic library link via DOI below!)

341. Mustafa SI, Schwarzacher T, and Heslop-Harrison JS. 2018. Complete mitogenomes from Kurdistani sheep: abundant centromeric nuclear copies representing diverse ancestorsMitochondrial DNA Part A () Mustafa et al_2018 AuthorVersion


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Morphology, adaptation and speciation

Plant adaptation arises from their morphology, itself a product of evolution and development. In this figure, the aspects and interactions of research at different levels are shown, with the work having implications across botany, including understanding plant phylogeny and speciation, and for ecology and ecosystems.

Plant adaptation arises from their morphology, itself a product of evolution and development. In this figure, the aspects and interactions of research at different levels are shown, with the work having implications across botany, including understanding plant phylogeny and speciation, and for ecology and ecosystems.

Heslop-Harrison JS. 2017. Morphology, adaptation and speciation. Annals of Botany 120(7): 621-624. study of plant evolution and development in a phylogenetic context has accelerated research advances in both areas over the last decade. The addition of a robust phylogeny for plant taxa based on DNA as well as morphology has given a strong context for this research. Genetics and genomics, including sequencing of many genes, and a better understanding of non-genetic, responsive changes, by plants have increased knowledge of how the different body forms of plants have arisen. Here, I overview the papers in this Special Issue of Annals of Botany on Morphological Adaptation, bringing together a range of papers that link phylogeny and morphology. These lead to models of development and functional adaptation across a range of plant systems, with implications for ecology and ecosystems, as well as development and evolution.

The study of evolution and development (evo-devo) has advanced plant research including speciation (Fernández-Mazuecos and Glover 2017); and as Theodore Dobzhansky stated in 1973, ‘nothing makes sense except in the light of evolution’. But what options are there for plant architecture to evolve – from the gene to cellular to organ and on to whole plant and ecosystem level? As suggested in the mind-map of Fig. 1, we are now in a good position to exploit data from morphological and genetic studies to understand the key processes of evolution and development, taking those results to find their impact on ecology and ecosystems. The papers in this special issue cover a diverse range of species, organs (related to leaves, roots and flowers) and approaches (from advanced microscopy to DNA fingerprinting), to show how modern plant studies can be integrated to lead to models of evolution and understand plant development in the broadest context.


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