J. Richard McIntosh

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J. Richard McIntosh is a Distinguished Professor Emeritus in Molecular, Cellular, and Developmental Biology at the University of Colorado Boulder. [1] McIntosh first graduated from Harvard with a BA in Physics in 1961, and again with a Ph.D. in Biophysics in 1968. [1] He began his teaching career at Harvard but has spent most of his career at the University of Colorado Boulder. [2] At the University of Colorado Boulder, McIntosh taught biology courses at both the undergraduate and graduate levels. [3] Additionally, he created an undergraduate course in the biology of cancer towards the last several years of his teaching career. [4] McIntosh's research career looks at a variety of things, including different parts of mitosis, microtubules, and motor proteins.

Contents

Research interests

Sequence of events in mitosis Mitosis cells sequence.svg
Sequence of events in mitosis

Mitosis

Most of McIntosh’s work focuses on the process of mitosis in the cell. Mitosis is the process of cell division that includes distinct movements of chromosomes in the cell and formation of mitotic spindles. [5] Additionally, McIntosh is very interested in the role of microtubules and motor proteins in this process. Mitotic spindles, composed of microtubules and other proteins, ensure that each of the two new cells during cell division both get one copy of every chromosome. [6] After the chromosomes are separated, then the cells are able to completely separate as well through cytokinesis. [6] In mitosis, there are multiple phases. [6] In prophase, the DNA starts to package itself for division and microtubules reorganize to prepare to form the mitotic spindle. [6] In prometaphase, kinetochores develop where the chromosome will attach to the mitotic spindle. [6] After that, chromosomes move towards the middle of the cell (metaphase plate) and the two copies separate during anaphase. [6] In some of McIntosh’s work he looks specifically at the anaphase A portion, which is related to where the chromosome is in relation to the pole it is moving towards. [6] Lastly, in telophase, the cell is wrapping up the stages of mitosis with creating a new nuclear envelope. [6]

Use of electron tomography

Additionally, in many of the works outlined below, McIntosh commonly uses electron tomography to image and study the cells. In electron tomography, many different images are put together to create a 3-D image of the subject being studied. [7] This technology is best suited to observe extremely complex structures and can image thinner sections of samples than can be physically made to study. [7]

1970s and 1980s

One of McIntosh’s earlier studies in the field of cell biology is in 1974, where his team published a paper on the structures of flagella of Pyrsonympha, an organism found in termites. [8] In this work, his team described the axostyle, a collection of microtubules, and presented that the axostyle’s attachment to other parts of the cell controls its function. [8]

In 1980, McIntosh’s curiosity with microtubules continued in “Visualization of the structural polarity of microtubules". [9] The polarity of microtubules is essential to generate the force needed to separate the chromosomes during mitosis, but at the time it was difficult to determine what the polarity is. [9] McIntosh’s team uses basal bodies and HeLa cells to study how protofilaments ‘hook’ onto them—either in right-handedness or left-handedness—invitro to determine polarity. [9]

A few years later, McIntosh published a study in 1984 on how tubulin moves in mammalian cells with a focus on the cell cycle. [10] To study the movements in tubulin in cells during mitosis and interphase, McIntosh used two imaging methods: labeled (dichlorotriazinyl-aminofluorescein or DTAF-) tubulin and fluorescence redistribution (or recovery) after photobleaching (or FRAP). [10] Using the labeled tubulin, McIntosh observed how quickly the freely-added labeled tubulin was polymerized to the existing microtubule structures in the cell. [10] It was noted that measuring the tubulin addition in interphase was difficult due to the lack of structures, while it was more observable in a mitotic cell. [10] While using FRAP, McIntosh noticed that the tubulin redistributed throughout the cytoplasm in both a rapid phase as well as a slower phase. [10] Overall, the redistribution or movement of the labeled tubulin in cells undergoing mitosis was much faster than the redistribution observed for cells in interphase. [10] The next year, McIntosh’s interests started to shift towards motor proteins. Kinesin, a motor protein found to move around vesicles in the cell, was recently discovered on another paper published the same year. [11] Here, McIntosh explored the possibility of kinesin as an important part of mitosis, as it can be found in the mitotic spindle. [11] Some of the possible functions that McIntosh suggested kinesin may have in mitosis are that they move chromosomes down microtubules or move microtubules in different areas within the mitotic spindle. [11]

The next year, McIntosh’s interests started to shift towards motor proteins. Kinesin, a motor protein found to move around vesicles in the cell, was recently discovered on another paper published the same year. [11] Here, McIntosh explored the possibility of kinesin as an important part of mitosis, as it can be found in the mitotic spindle. [11] Some of the possible functions that McIntosh suggested kinesin may have in mitosis are that they move chromosomes down microtubules or move microtubules in different areas within the mitotic spindle. [11]

1990s

McIntosh’s interest in the connection of motor proteins and mitosis continues into the 1990s. In 1990, he published a paper on cytoplasmic dynein and mitosis. [12] Dynein, a protein first found in the flagella, had now been discovered in the cytoplasm as well. [12] It moves towards the minus end, or the slower-growing end, of the microtubules. [12] The research team used antibodies to image the distribution of cytoplasmic dynein in the cell. [12] Dynein was found to be near kinetochores during mitosis, while in interphase they were scattered around the cell. [12] The research team used this finding to suggest that dynein had a role in how chromosomes separate during mitosis due to the spatial differences of dynein during interphase and mitosis. [12]

In contrast to the topics McIntosh had researched so far, in 1996 his team published “Computer Visualization of Three-Dimensional Image Data using IMOD.” [13] In this article, his team shared the development of the computer software called IMOD. [13] With IMOD, researchers can study tomographies and data from both electron and light microscopes, and create three-dimensional images to interact with. [13] With these reconstructions, IMOD gives researchers the ability to help visualize their samples digitally. [7] The software is able to create collections of images and uses these collections to make models and measurements for further analysis. [7] One tool, the ‘slicer’, can help view samples at different angles. [7] Some other tools include zooming and panning (called the Zap window), a Model view window of a contour of the sample, an XYZ window that shows the other planes that cut through the point that is being studied, and the Tilt and Tumbler windows that show different projections that can be made. [13] The software can be downloaded and used for free at http://bio3d.colorado.edu. [7] This technology has been used to visualize cytoplasmic membranes, myofibrils, and the trans-Golgi network. [13]

A few years later in 1999, McIntosh’s team published a study using cryofixing and electron tomography to create a 3-dimensional model of the Golgi apparatus. [14] Two of the Golgi apparatus’s main functions are to modify proteins and to target them to their next destination. [14] For transport within the Golgi, McIntosh’s team proposed evidence that it can be done by vesicles in the cell that fuse with different membranes or by microtubules that are constantly forming and shifting around the Golgi. [14] By cryofixing, or rapidly freezing the cells to be studied, McIntosh’s team was able to preserve the cells essentially in time before using an electron microscope to visualize them. [14] In this work, the researchers visualized different coatings on the budding parts of the Golgi. [14] They were able to see the difference between clathrin-coated and non-clathrin-coated vesicles. [14] The buds help molecules in the Golgi be transported from the Golgi to their intended area in the cell. [14] In this work, the researchers describe the different cisternae, buds, vesicles, and coatings and the differences between the cis and trans faces of the Golgi. [14] This visualization technique led them to the conclusion that the different cisternae of the Golgi are not interconnected. [14] There are also many vesicles that surround the Golgi, most of which are not coated with clathrin. [14]

2000s

In 2002, McIntosh’s team continued his earlier interests in “Chromosome-microtubule interactions during mitosis.” [15] This review paper explains how spindle microtubules bind to chromosomes during segregation at places called kinetochores and the description of the Kinetochore-Dependent Checkpoint. [15] This point in the cell will block the segregation of chromosomes until all of the chromosomes are properly connected. [15] It also explains the role of microtubule proteins, motor proteins, and the microtubules themselves in the segregation process. [15] Some of the motor proteins mentioned in the article include those associated with the kinetochore at the plus or minus end or those that help with the disassembly of microtubules. [15] Some motor proteins associate with the chromosomes instead of the kinetochores, including chromokinesins, and some work at the spindle poles. [15]

McIntosh also contributed to a work published in 2004 by many scientists in “A standard kinesin nomenclature”. [16] In this work, these scientists helped create a naming structure for kinesin proteins, that are involved in transport in the cell along with microtubules. [16] The kinesin proteins were split into fourteen different families, and given different names such as kinesin-1, kinesin-2, etc. [16] This new structure of naming was created so that the existing kinesins can be easily classified as well as classify new kinesin proteins when they are discovered. [16] The authors recommend using protein sequence alignment based homology search to help classify the kinesins. [16] It also makes specifications on what makes a kinesin recognized. [16] Additionally, this paper also gives guidance on how to address kinesins (and their former names, if applicable) in papers. [16]

In 2005, McIntosh and his team published a study that investigated the link between the depolymerization of microtubules and the generation of force, showing how the chemistry of the tubulin dimers can create mechanical force. [17] The researchers attached glass microbeads to tubulin. [17] By using laser tweezers, McIntosh’s team was able to observe that the bead would move, typically towards the minus end, as the microtubules depolymerized off of it. [17] In particular, the bending of the protofilaments would cause the tweezers to pick up the force generated. [17] Here, the researchers applied these findings to chromosome segregation and concluded that the microtubule dynamics create the forces needed during mitosis. [17]

In 2006, McIntosh’s team used cyroelecton tomography to image axonemes in sea urchin sperm and Chlamydomonas reinhardtii. [18] Axonemes are structures in cilia and flagella consisting of a specific pattern of microtubules. [18] McIntosh’s team was particularly interested in the dynein protein. [18] In this paper, the researchers describe how the different subunits of dynein attach to the microtubules and how they might generate force as well. [18]

2010s

McIntosh’s letter “Motors or dynamics: What really moves chromosomes?” to Nature cell biology in 2012 explains an overview of the different directions his lab has taken so far. [19] In those who study how chromosomes move during the process of mitosis, researchers argue either that microtubules generate the forces needed to separate them or that certain motor proteins do. [19] When McIntosh first started his research, he noted he was very confident in the motor protein school of thought and related it to the sliding filament theory used to explain muscle contraction. [19] However, by reading other research group’s papers and conducting other studies that found adenosine triphosphate (ATP) and the motor protein dynein were not necessary for chromosome movement, he started to consider the influence that microtubules had in this process. [19]

Continuing on the interest of microtubules, the mitotic spindle, and mitosis, in 2013 his team published a paper studying the speed of chromosome movement during mitosis. [20] It was observed overall that microtubule depolymerization and the movement of motor proteins are very fast, but the chromosomes move slowly during their separation in the anaphase A stage of mitosis. [20] In this study, McIntosh and his team suggest that motor proteins could have an influence in the rate of movement by influencing depolymerization. [20] It was also noted that separation of different protofilaments over individual tubulin dimers may be another influence in the depolymerization and subsequent chromosome movement in mitosis. [20]

Focusing now on microtubule polymerization instead of depolymerization, in 2018 McIntosh’s team used electron tomography to study the microtubules in vitro as well as in six different species. [21] The researchers noticed as the microtubules grow, they bend out from the axis, similar to how the microtubules bend off when they are depolymerizing as well. [21]

2020s

Continuing using the electron tomography technique, in 2020 McIntosh and his team used this visualization strategy to observe how different kinds of microtubules work together during the metaphase state of mitosis. [22] One important structure in this phase is kinetochore microtubules or referred to as KMTs. [22] They, along with other structures in the cell, work together to balance out the different forces in the cell during mitosis. [22]

Books

Throughout his career, McIntosh has helped edit and write books as well. In 2001, he was an editor along with Joseph G. Gall (from the Carnegie Institution of Washington) in Landmark Papers in Cell Biology. [23] This work, celebrating the 40th anniversary of the creation of the American Society for Cell Biology (founded in 1960) includes 42 major papers in the field of cell biology along with the editor’s own thoughts. [23] Some of the topics in this book include transcription, mitosis, cell membrane, and the cytoskeleton. [23]

In 2017, McIntosh is the guest editor of Mechanisms of Mitotic Chromosome Segregation. [24] In this work, McIntosh connects various findings about mitosis from a variety of organisms. [24] In the introduction, McIntosh also recognizes the importance of how the evolution of our understanding of mitosis comes from the advancement of other technologies in the field such as better cameras, better ways to purify molecules, and an increase in the understanding of genetics. [24] This collection of review articles helps readers get an overview of mitosis, a process that McIntosh believes is essential to life itself. [24]

Published in 2019, Understanding Cancer: An Introduction to the Biology, Medicine, and Societal Implications of the Disease is a resource for those who want to learn holistically more about cancer. [4] This book, influenced by his son’s death to lung cancer, discusses every stage of the process: screening, diagnosing, and treating. [4] The text is written so that those without previous knowledge are able to learn the basics about cancer. [4] McIntosh recognizes in the introduction that cancer is just more than science; it encompasses other fields such as history and religion. [2] In this text, McIntosh discusses science-heavy topics related to cancer such as the role of oncogenes, tumor suppressors, and the immune system. [4] Additionally, the text covers topics beyond the scientific descriptions of cancer such as on the future of cancer, minimizing the risk of cancer, and living with cancer. [4]

Online lectures

Recorded in late 2008, McIntosh is featured on the website “iBiology” giving a series of talks titled “Eukaryotic cell division.” [2] In this series, the process is split up into three different videos specializing in chromosome division, experimentation, and the mitotic stage of anaphase A. [2]

Accolades

From 1984 to 2006, McIntosh served as the Director of the Boulder Laboratory for 3-D Electron Microscopy of Cells. [1] In 1994, McIntosh served as a Research Professor of the American Cancer Society and continued this role until 2006. [1] In 1994, McIntosh was also the President of the American Society for Cell Biology. [1] In 1999, McIntosh was elected to both the National Academy of Sciences and the American Academy of Arts and Sciences. [1] The National Academy of Sciences lists him as a primary member of the Cellular and Developmental Biology section and a secondary member of the Biophysics and Computational Biology section. [25] In 2000, McIntosh was award with the Distinguished Professor title at the University of Colorado, Boulder. [1] After he retired in 2006, McIntosh continues to conduct research and publish books and articles. [23]

Related Research Articles

<span class="mw-page-title-main">Mitosis</span> Process in which replicated chromosomes are separated into two new identical nuclei

In cell biology, mitosis is a part of the cell cycle in which replicated chromosomes are separated into two new nuclei. Cell division by mitosis gives rise to genetically identical cells in which the total number of chromosomes is maintained. Therefore, mitosis is also known as equational division. In general, mitosis is preceded by S phase of interphase and is often followed by telophase and cytokinesis; which divides the cytoplasm, organelles and cell membrane of one cell into two new cells containing roughly equal shares of these cellular components. The different stages of mitosis altogether define the mitotic (M) phase of a cell cycle—the division of the mother cell into two daughter cells genetically identical to each other.

<span class="mw-page-title-main">Microtubule</span> Polymer of tubulin that forms part of the cytoskeleton

Microtubules are polymers of tubulin that form part of the cytoskeleton and provide structure and shape to eukaryotic cells. Microtubules can be as long as 50 micrometres, as wide as 23 to 27 nm and have an inner diameter between 11 and 15 nm. They are formed by the polymerization of a dimer of two globular proteins, alpha and beta tubulin into protofilaments that can then associate laterally to form a hollow tube, the microtubule. The most common form of a microtubule consists of 13 protofilaments in the tubular arrangement.

<span class="mw-page-title-main">Cell division</span> Process by which living cells divide

Cell division is the process by which a parent cell divides into two daughter cells. Cell division usually occurs as part of a larger cell cycle in which the cell grows and replicates its chromosome(s) before dividing. In eukaryotes, there are two distinct types of cell division: a vegetative division (mitosis), producing daughter cells genetically identical to the parent cell, and a cell division that produces haploid gametes for sexual reproduction (meiosis), reducing the number of chromosomes from two of each type in the diploid parent cell to one of each type in the daughter cells. In cell biology, mitosis (/maɪˈtoʊsɪs/) is a part of the cell cycle, in which, replicated chromosomes are separated into two new nuclei. Cell division gives rise to genetically identical cells in which the total number of chromosomes is maintained. In general, mitosis is preceded by the S stage of interphase and is often followed by telophase and cytokinesis; which divides the cytoplasm, organelles, and cell membrane of one cell into two new cells containing roughly equal shares of these cellular components. The different stages of mitosis all together define the mitotic (M) phase of animal cell cycle—the division of the mother cell into two genetically identical daughter cells. Meiosis undergoes two divisions resulting in four haploid daughter cells. Homologous chromosomes are separated in the first division of meiosis, such that each daughter cell has one copy of each chromosome. These chromosomes have already been replicated and have two sister chromatids which are then separated during the second division of meiosis. Both of these cell division cycles are used in the process of sexual reproduction at some point in their life cycle. Both are believed to be present in the last eukaryotic common ancestor.

<span class="mw-page-title-main">Anaphase</span> Stage of a cell division

Anaphase is the stage of mitosis after the process of metaphase, when replicated chromosomes are split and the newly-copied chromosomes are moved to opposite poles of the cell. Chromosomes also reach their overall maximum condensation in late anaphase, to help chromosome segregation and the re-formation of the nucleus.

<span class="mw-page-title-main">Spindle apparatus</span> Feature of biological cell structure

In cell biology, the spindle apparatus is the cytoskeletal structure of eukaryotic cells that forms during cell division to separate sister chromatids between daughter cells. It is referred to as the mitotic spindle during mitosis, a process that produces genetically identical daughter cells, or the meiotic spindle during meiosis, a process that produces gametes with half the number of chromosomes of the parent cell.

<span class="mw-page-title-main">Telophase</span> Final stage of a cell division for eukaryotic cells both in mitosis and meiosis

Telophase is the final stage in both meiosis and mitosis in a eukaryotic cell. During telophase, the effects of prophase and prometaphase are reversed. As chromosomes reach the cell poles, a nuclear envelope is re-assembled around each set of chromatids, the nucleoli reappear, and chromosomes begin to decondense back into the expanded chromatin that is present during interphase. The mitotic spindle is disassembled and remaining spindle microtubules are depolymerized. Telophase accounts for approximately 2% of the cell cycle's duration.

The microtubule-organizing Centre (MTOC) is a structure found in eukaryotic cells from which microtubules emerge. MTOCs have two main functions: the organization of eukaryotic flagella and cilia and the organization of the mitotic and meiotic spindle apparatus, which separate the chromosomes during cell division. The MTOC is a major site of microtubule nucleation and can be visualized in cells by immunohistochemical detection of γ-tubulin. The morphological characteristics of MTOCs vary between the different phyla and kingdoms. In animals, the two most important types of MTOCs are 1) the basal bodies associated with cilia and flagella and 2) the centrosome associated with spindle formation.

<span class="mw-page-title-main">Dynein</span> Class of enzymes

Dyneins are a family of cytoskeletal motor proteins that move along microtubules in cells. They convert the chemical energy stored in ATP to mechanical work. Dynein transports various cellular cargos, provides forces and displacements important in mitosis, and drives the beat of eukaryotic cilia and flagella. All of these functions rely on dynein's ability to move towards the minus-end of the microtubules, known as retrograde transport; thus, they are called "minus-end directed motors". In contrast, most kinesin motor proteins move toward the microtubules' plus-end, in what is called anterograde transport.

<span class="mw-page-title-main">Spindle checkpoint</span> Cell cycle checkpoint

The spindle checkpoint, also known as the metaphase-to-anaphase transition, the spindle assembly checkpoint (SAC), the metaphase checkpoint, or the mitotic checkpoint, is a cell cycle checkpoint during metaphase of mitosis or meiosis that prevents the separation of the duplicated chromosomes (anaphase) until each chromosome is properly attached to the spindle. To achieve proper segregation, the two kinetochores on the sister chromatids must be attached to opposite spindle poles. Only this pattern of attachment will ensure that each daughter cell receives one copy of the chromosome. The defining biochemical feature of this checkpoint is the stimulation of the anaphase-promoting complex by M-phase cyclin-CDK complexes, which in turn causes the proteolytic destruction of cyclins and proteins that hold the sister chromatids together.

<span class="mw-page-title-main">Kinetochore</span> Protein complex that allows microtubules to attach to chromosomes during cell division

A kinetochore is a disc-shaped protein structure associated with duplicated chromatids in eukaryotic cells where the spindle fibers attach during cell division to pull sister chromatids apart. The kinetochore assembles on the centromere and links the chromosome to microtubule polymers from the mitotic spindle during mitosis and meiosis. The term kinetochore was first used in a footnote in a 1934 Cytology book by Lester W. Sharp and commonly accepted in 1936. Sharp's footnote reads: "The convenient term kinetochore has been suggested to the author by J. A. Moore", likely referring to John Alexander Moore who had joined Columbia University as a freshman in 1932.

<span class="mw-page-title-main">Motor protein</span> Class of molecular proteins

Motor proteins are a class of molecular motors that can move along the cytoplasm of cells. They convert chemical energy into mechanical work by the hydrolysis of ATP. Flagellar rotation, however, is powered by a proton pump.

A spindle poison, also known as a spindle toxin, is a poison that disrupts cell division by affecting the protein threads that connect the centromere regions of chromosomes, known as spindles. Spindle poisons effectively cease the production of new cells by interrupting the mitosis phase of cell division at the spindle assembly checkpoint (SAC). However, as numerous and varied as they are, spindle poisons are not yet 100% effective at ending the formation of tumors (neoplasms). Although not 100% effective, substantive therapeutic efficacy has been found in these types of chemotherapeutic treatments. The mitotic spindle is composed of microtubules that aid, along with regulatory proteins, each other in the activity of appropriately segregating replicated chromosomes. Certain compounds affecting the mitotic spindle have proven highly effective against solid tumors and hematological malignancies.

KRP stands for kinesin related proteins. bimC is a subfamily of KRPs and its function is to separate the duplicated centrosomes during mitosis.

The Kinesin-13 Family are a subfamily of motor proteins known as kinesins. Most kinesins transport materials or cargo around the cell while traversing along microtubule polymer tracks with the help of ATP-hydrolysis-created energy.

<span class="mw-page-title-main">Aurora kinase B</span> Protein

Aurora kinase B is a protein that functions in the attachment of the mitotic spindle to the centromere.

<span class="mw-page-title-main">Dynactin</span>

Dynactin is a 23 subunit protein complex that acts as a co-factor for the microtubule motor cytoplasmic dynein-1. It is built around a short filament of actin related protein-1 (Arp1).

The Kinesin 8 Family are a subfamily of the molecular motor proteins known as kinesins. Most kinesins transport materials or cargo around the cell while traversing along microtubule polymer tracks with the help of ATP-hydrolysis-created energy. The Kinesin 8 family has been shown to play an important role in chromosome alignment during mitosis. Kinesin 8 family members KIF18A in humans and Kip3 in yeast have been shown to be in vivo plus-end directed microtubule depolymerizers. During prometaphase of mitosis, the microtubules attach to the kinetochores of sister chromatids. Kinesin 8 is thought to play some role in this process, as knockdown of this protein via siRNA produces a phenotype of sister chromatids that are unable to align properly.

Conly Leroy Rieder is a cancer researcher in the field of mitotic cellular division. The bulk of his research between 1980 and 2011 was funded through NIH grants and conducted at the Wadsworth Center in the New York State Department of Health in Albany, New York. He has published on the subjects of chromosome motility, spindle assembly, and mitotic checkpoints. His research has contributed to the growing understanding of the process of cell division and the pathology of cancer.

<span class="mw-page-title-main">Microtubule plus-end tracking protein</span>

Microtubule plus-end/positive-end tracking proteins or +TIPs are a type of microtubule associated protein (MAP) which accumulate at the plus ends of microtubules. +TIPs are arranged in diverse groups which are classified based on their structural components; however, all classifications are distinguished by their specific accumulation at the plus end of microtubules and their ability to maintain interactions between themselves and other +TIPs regardless of type. +TIPs can be either membrane bound or cytoplasmic, depending on the type of +TIPs. Most +TIPs track the ends of extending microtubules in a non-autonomous manner.

<span class="mw-page-title-main">Neurotubule</span>

Neurotubules are microtubules found in neurons in nervous tissues. Along with neurofilaments and microfilaments, they form the cytoskeleton of neurons. Neurotubules are undivided hollow cylinders that are made up of tubulin protein polymers and arrays parallel to the plasma membrane in neurons. Neurotubules have an outer diameter of about 23 nm and an inner diameter, also known as the central core, of about 12 nm. The wall of the neurotubules is about 5 nm in width. There is a non-opaque clear zone surrounding the neurotubule and it is about 40 nm in diameter. Like microtubules, neurotubules are greatly dynamic and the length of them can be adjusted by polymerization and depolymerization of tubulin.

References

  1. 1 2 3 4 5 6 7 "J. Richard McIntosh". Biophysics Group. 2015-11-13. Retrieved 2021-04-15.
  2. 1 2 3 4 "Richard 'Dick' McIntosh • iBiology". iBiology. Retrieved 2021-04-15.
  3. ↑ "Richard McIntosh - Routledge & CRC Press Author Profile". www.routledge.com. Retrieved 2021-04-15.
  4. 1 2 3 4 5 6 McIntosh JR (2019). Understanding cancer : an introduction to the biology, medicine, and societal implications of this disease. Boca Raton. ISBN   978-0-8153-4535-0. OCLC   1066186470.{{cite book}}: CS1 maint: location missing publisher (link)
  5. ↑ McIntosh JR, Hays T (December 2016). "A Brief History of Research on Mitotic Mechanisms". Biology. 5 (4): 55. doi: 10.3390/biology5040055 . PMC   5192435 . PMID   28009830.
  6. 1 2 3 4 5 6 7 8 McIntosh JR (September 2016). "Mitosis". Cold Spring Harbor Perspectives in Biology. 8 (9): a023218. doi:10.1101/cshperspect.a023218. PMC   5008068 . PMID   27587616.
  7. 1 2 3 4 5 6 O'Toole ET, Winey M, McIntosh JR, Mastronarde DN (2002-01-01). "Electron tomography of yeast cells". Guide to Yeast Genetics and Molecular and Cell Biology Part C. Methods in Enzymology. Vol. 351. pp. 81–95. doi:10.1016/S0076-6879(02)51842-5. ISBN   9780121822545. PMC   4440791 . PMID   12073377.
  8. 1 2 Bloodgood RA, Miller KR, Fitzharris TP, Mcintosh JR (May 1974). "The ultrastructure of Pyrsonympha and its associated microorganisms". Journal of Morphology. 143 (1): 77–105. doi:10.1002/jmor.1051430104. PMID   30326674. S2CID   53528610.
  9. 1 2 3 Heidemann SR, McIntosh JR (July 1980). "Visualization of the structural polarity of microtubules". Nature. 286 (5772): 517–9. Bibcode:1980Natur.286..517H. doi:10.1038/286517a0. PMID   7402333. S2CID   1656543.
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  11. 1 2 3 4 5 6 Scholey JM, Porter ME, Grissom PM, McIntosh JR (December 1985). "Identification of kinesin in sea urchin eggs, and evidence for its localization in the mitotic spindle". Nature. 318 (6045): 483–6. Bibcode:1985Natur.318..483S. doi:10.1038/318483a0. PMID   2933590. S2CID   4345279.
  12. 1 2 3 4 5 6 Pfarr CM, Coue M, Grissom PM, Hays TS, Porter ME, McIntosh JR (May 1990). "Cytoplasmic dynein is localized to kinetochores during mitosis". Nature. 345 (6272): 263–5. Bibcode:1990Natur.345..263P. doi:10.1038/345263a0. PMID   2139717. S2CID   4364671.
  13. 1 2 3 4 5 Kremer JR, Mastronarde DN, McIntosh JR (1996-01-01). "Computer visualization of three-dimensional image data using IMOD". Journal of Structural Biology. 116 (1): 71–6. doi:10.1006/jsbi.1996.0013. PMID   8742726. S2CID   4608130.
  14. 1 2 3 4 5 6 7 8 9 10 Ladinsky MS, Mastronarde DN, McIntosh JR, Howell KE, Staehelin LA (March 1999). "Golgi structure in three dimensions: functional insights from the normal rat kidney cell". The Journal of Cell Biology. 144 (6): 1135–49. doi:10.1083/jcb.144.6.1135. PMC   2150572 . PMID   10087259.
  15. 1 2 3 4 5 6 McIntosh JR, Grishchuk EL, West RR (2002-11-01). "Chromosome-microtubule interactions during mitosis". Annual Review of Cell and Developmental Biology. 18 (1): 193–219. doi:10.1146/annurev.cellbio.18.032002.132412. PMID   12142285.
  16. 1 2 3 4 5 6 7 Lawrence CJ, Dawe RK, Christie KR, Cleveland DW, Dawson SC, Endow SA, et al. (October 2004). "A standardized kinesin nomenclature". The Journal of Cell Biology. 167 (1): 19–22. doi:10.1083/jcb.200408113. PMC   2041940 . PMID   15479732.
  17. 1 2 3 4 5 Grishchuk EL, Molodtsov MI, Ataullakhanov FI, McIntosh JR (November 2005). "Force production by disassembling microtubules". Nature. 438 (7066): 384–8. Bibcode:2005Natur.438..384G. doi:10.1038/nature04132. PMID   16292315. S2CID   4415883.
  18. 1 2 3 4 Nicastro D, Schwartz C, Pierson J, Gaudette R, Porter ME, McIntosh JR (August 2006). "The molecular architecture of axonemes revealed by cryoelectron tomography". Science. 313 (5789): 944–8. Bibcode:2006Sci...313..944N. doi:10.1126/science.1128618. PMID   16917055. S2CID   43436284.
  19. 1 2 3 4 McIntosh JR (December 2012). "Motors or dynamics: what really moves chromosomes?". Nature Cell Biology. 14 (12): 1234. doi:10.1038/ncb2649. PMC   4429872 . PMID   23196840.
  20. 1 2 3 4 Betterton MD, McIntosh JR (December 2013). "Regulation of chromosome speeds in mitosis". Cellular and Molecular Bioengineering. 6 (4): 418–430. doi:10.1007/s12195-013-0297-4. PMC   4578309 . PMID   26405462.
  21. 1 2 McIntosh JR, O'Toole E, Morgan G, Austin J, Ulyanov E, Ataullakhanov F, Gudimchuk N (August 2018). "Microtubules grow by the addition of bent guanosine triphosphate tubulin to the tips of curved protofilaments". The Journal of Cell Biology. 217 (8): 2691–2708. doi:10.1083/jcb.201802138. PMC   6080942 . PMID   29794031.
  22. 1 2 3 O'Toole E, Morphew M, McIntosh JR (February 2020). "Electron tomography reveals aspects of spindle structure important for mechanical stability at metaphase". Molecular Biology of the Cell. 31 (3): 184–195. doi:10.1091/mbc.E19-07-0405. PMC   7001478 . PMID   31825721.
  23. 1 2 3 4 Gall JG (November 2000). Landmark Papers in Cell Biology : Selected Research Articles Celebrating Forty Years of The ASCB. J. Richard McIntosh. Plymouth: Cold Spring Harbor Laboratory Press. ISBN   0-87969-602-8. OCLC   697711820.
  24. 1 2 3 4 Mechanisms of mitotic chromosome segregation. [Place of publication not identified]: MDPI AG. 2017. ISBN   978-3-03842-402-4. OCLC   990847914.
  25. ↑ "J. Richard McIntosh". www.nasonline.org. Retrieved 2021-04-15.
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