Peter K. Hepler









































Peter K. Hepler

Peter Hepler.jpg
Born
October 29, 1936
Dover, New Hampshire
Nationality
American
Citizenship
United States
Alma mater
University of New Hampshire, B.S. Chemistry 1958
University of Wisconsin, Ph.D. Plant Cell Biology 1964
Known for
Cell biology, plant physiology, microscopy

Scientific career
Fields
Cell biology, plant physiology, microscopy
Institutions
Stanford University
University of Massachusetts at Amherst

Website

Peter K. Hepler


Molecular & Cellular Biology

Peter Klock Hepler HonFRMS is the Constantine J. Gilgut and Ray Ethan Torrey Professor Emeritus in the Biology Department of the University of Massachusetts at Amherst who is notable for his work on elucidating the roles of calcium,[1]membranes[2] and the cytoskeleton[3][4] in plant cell development and cell motility.




Contents






  • 1 Personal life


  • 2 University life


  • 3 Research


    • 3.1 Microtubules and cell shape


    • 3.2 Microtubules and cell motility


    • 3.3 Microfilaments and cytoplasmic streaming


    • 3.4 Calcium and plant development




  • 4 Honors and awards


  • 5 References





Personal life


Peter Klock Hepler was born on October 29, 1936, in Dover, New Hampshire, to Jesse Raymond Hepler[5][6][7] and Rebecca Orpha Peterson Hepler. He married Margaret (Peggy) Dennison Hunt on March 7, 1964. They have three children: Sarah, Lukas and Anna.[8] In an interview published in the Newsletter of the American Society of Plant Biologists, Hepler was asked, "What is your most treasured possession?" He answered, "My family; but I don't possess them."[9] Peter and Peggy Hepler live on a farm in Pelham, Massachusetts that was established by John Gray in 1740[10] and is now a part of the Kestrel Land Trust.[11]



University life


Peter Hepler graduated from Dover High School in 1954. He received his B.S. in chemistry from the University of New Hampshire in 1958 and earned his Ph.D. in plant cell biology from University of Wisconsin in 1964, studying the role of cortical microtubules in plant cell development with Eldon H. Newcomb. After receiving his Ph.D., Hepler served at the Walter Reed Army Institute of Research until 1966, studying malarial parasites. Hepler then returned to the University of Wisconsin for a postdoctoral fellowship[12] and then became a postdoctoral fellow with Keith Porter[13] at Harvard University from 1966-1967, where he continued his investigation of microtubules, focusing on their role in the mitotic apparatus and the phragmoplast of the endosperm cells of Haemanthus Katharinae. After being an assistant professor at Stanford University, Hepler joined the faculty in the Botany Department at the University of Massachusetts at Amherst. He was an associate professor from 1977 to 1980, a professor from 1980-1989, and became the Ray Ethan Torrey Professor in 1989 and the Constantine J. Gilgut Professor in 1998. Hepler retired from the Biology Department as the Constantine J. Gilgut and Ray Ethan Torrey Professor Emeritus, although he continues to do research.[14] Hepler spent many summers teaching and doing research at the Marine Biological Laboratory[15][16] at Woods Hole, Massachusetts. Hepler also participated in a multiyear international collaboration with Brian E. S. Gunning.[17]


Hepler was an Associate Editor of Protoplasma from 1994-2001 and Associate Editor of Plant Physiology from 1998-2000. He has been on the editorial boards of the Annual Review Plant Physiology, Plant and Cell Physiology, the Journal of Submicroscopic Cytology, Cell Motility and the Cytoskeleton, and BioEssays.[citation needed]



Research


Hepler's scientific method is to know thoroughly the classical botanical literature and then develop or apply modern physico-chemical techniques to answer salient and extensive biological questions using plants that are well-suited to answer those questions. In so doing, Hepler opened whole areas of research.[18][19] Hepler did pioneering work in showing the relationship of the microscopic elements of the cytoskeleton to the macroscopic properties of plant growth, development and function. He also did pioneering work on plasmodesmata,[20][21][22]stomatal function,[23][24][25][26] and in the development of techniques useful for answering questions using light[27][28][29][30][31] and electron microscopy.[32] Hepler's scientific publications with Barry A. Palevitz are notable for quoting Woody Allen and Yogi Berra.[33]


Hepler described his realization of the influence a review he and Palevitz[4] wrote on microtubules and microfilaments "to introduce new thoughts and promising avenues for future research" had with his characteristic self-deprecating sense of humor: "I became aware that the review was being read widely one summer (1979) while working in the library at the Marine Biological Laboratory. I turned to the library's volume of the Annual Review of Plant Physiology that contained our paper and when I put the volume down, it literally fell open at our article; worn edges on the pages and the penciled corrections of all the misspellings and punctuation errors indicated that the chapter had been thoroughly perused."[4]


Hepler, along with Ledbetter and Porter,[34] is considered to be a co-discoverer of microtubules.[13]



Microtubules and cell shape


In late 1962 and early 1963, Hepler tested the newly developed procedure using a glutaraldehyde pre-fix followed by an osmium post-fix to study plant cell structure using an electron microscope.[35] Building on the earlier work by Sinnott and Bloch,[36] who had shown that wounding the existing tracheary elements in a Coleus stem induced neighboring parenchyma cells to differentiate into new tracheary elements, Hepler showed that cytoplasmic microtubules were localized specifically in the cortical cytoplasm immediately over the bands of new secondary wall thickenings.[37] Moreover, Hepler discovered that the microtubules were oriented parallel to the cellulose microfibrils of the newly formed secondary wall thickenings. This work, along with the studies of Ledbetter and Porter[34] and Green[38] established the importance of cortical microtubules in controlling the alignment of cellulose microfibrils in the cell wall.[39][40] Further work with Barry Palevitz showed that microtubules were involved in orienting the cellulose microfibrils in the walls of guard cells in a pattern of radial micellation that is necessary for stomatal function.[41] Hepler, along with the husband and wife team of Dale Callaham and Sue Lancelle, developed a method to achieve rapid freeze fixation of particularly small plant cells that showed that cortical microtubules are closely associated with one another, actin microfilaments, the endoplasmic reticulum and the plasma membrane.[32][42]



Microtubules and cell motility


Building on the work of Shinya Inoué and Andrew Bajer using polarized light microscopy,[43] Hepler used electron microscopy to elucidate the nature of the microtubule/chromosome attachments at the kinetochore as well as the arrangement of the microtubules in the phragmoplast during the development of the new cell wall, where microtubules from both sides of the phragmoplast were seen to overlap with one another in the plane of the cell plate.[44]


Hepler realized that microtubules were dynamic structures that were deployed in various locations throughout the cell, and became interested in the mechanisms involved in microtubule organization in cells that lacked a microtubule-organizing center known as the centrosome. In order to understand how microtubule-organizing centers were generated, Hepler examined the de novo formation of the blepharoplast in the spermatogenous cells of Marsilea vestita. The blepharoplast in each spermatid generates 100–150 basal bodies, each of which gives rise to the 9+2 arrangement of microtubules in a cilium. During telophase of the penultimate division, flocculent material appears near clefts on the distal surfaces of the daughter nuclei. During prophase of the final division which gives rise to the spermatids, the flocculent material near each nucleus condenses to give rise to two blepharoplasts, which then separate, one going to each spermatid.[45]


While Hepler was successful in identifying an aggregation of material that possessed microtubule-organizing capacity, he was not able to specify the biophysical mechanisms involved in organization. After Richard Weisenberg[46] discovered that microtubule polymerization was sensitive to calcium concentration, Hepler realized that he had already seen a close association between elements of the endoplasmic reticulum and microtubules in the mitotic apparatus and in the phragmoplast and suggested that these membranes may function in controlling the concentration of free calcium in the mitotic apparatus.[47] Along with Susan Wick and Steve Wolniak, Hepler showed that the endoplasmic reticulum contained stores of calcium and suggested that the endoplasmic reticulum may locally control the calcium concentration and thus the polymerization/depolymerization of microtubules. Subsequently,[48][49] Hepler, along with Dale Callaham, Dahong Zhang, and Patricia Wadsworth, observed calcium ion transients during mitosis[50][51] and showed that the microinjection of calcium ions into the mitotic spindle does regulate the depolymerization of microtubules and the movement of chromosomes to the poles during mitosis.[52][53][54]



Microfilaments and cytoplasmic streaming


Hepler identified actin microfilaments in bundles at the ectoplasm-endoplasm interface of Nitella internodal cells by showing that the bundles bound heavy meromyosin, giving the characteristic arrowhead arrangement.[55][56] The actin microfilaments had the correct polarity to be part of the actomyosin motor that provides the motive force for cytoplasmic streaming in these giant algal cells.[57]



Calcium and plant development


Hepler has shown that calcium ions are a central regulator of plant growth and development[58] specifically demonstrating that calcium is important for tip growth[59][60][61] and in phytochrome.[62][63] and cytokinin[64][65][66] action.



Honors and awards



  • In 1975, Hepler was the fourth recipient of the Jeanette Siron Pelton Award given by the Botanical Society of America, because his "penetrating analytical and experimental studies of the ultrastructure of differentiating cells have made a significant and lasting contribution to our perception of morphogenesis at the cellular level. In particular his work on the ultrastructure of differentiating xylem elements, on the roles of microtubules and microfibrils, and on the control of the orientation of mitotic spindles in differentiating cells have provided new insights which hold great promise for the future."[67]

  • In 2007, Hepler was named an inaugural Fellow of the American Society of Plant Biologists.[68]

  • In 2010, Hepler was elected as a Fellow of the American Association for the Advancement of Science for his contributions as "one of the most influential plant cell biologists, who has continuously and continues to achieve breakthroughs that have guided research directions of numerous plant scientists."[18][19][69]

  • In 2011, Hepler was honored with the Charles Reid Barnes Life Membership Award from the American Society of Plant Biologists.[70]

  • In 2015, Hepler was named an Honorary Fellow of the Royal Microscopical Society for his contributions to plant science, including publishing the first report suggesting a co-alignment of microtubules with cell wall cellulose microtubules.[71][72]

  • A scholarship was named in honor of Hepler. The Peter K. Hepler Research Scholarship supports undergraduate research on a biological question in a laboratory or field setting outside of the United States.[73]

  • The Plant Biology Graduate Program at the University of Massachusetts Amherst held a symposium on October 14, 2017 entitled: Capturing the dynamic architecture of cells: Honoring the high-resolution career of Peter Hepler. Friends, family, students, and colleagues celebrated his life and contributions to plant cell biology.[74]



References





  1. ^ Hepler, P. K.; R. O. Wayne (1993). "This Week's Citation Classic" (PDF). Current Contents (July 26, 1993) (30): 8. Retrieved October 6, 2016..mw-parser-output cite.citation{font-style:inherit}.mw-parser-output q{quotes:"""""""'""'"}.mw-parser-output code.cs1-code{color:inherit;background:inherit;border:inherit;padding:inherit}.mw-parser-output .cs1-lock-free a{background:url("//upload.wikimedia.org/wikipedia/commons/thumb/6/65/Lock-green.svg/9px-Lock-green.svg.png")no-repeat;background-position:right .1em center}.mw-parser-output .cs1-lock-limited a,.mw-parser-output .cs1-lock-registration a{background:url("//upload.wikimedia.org/wikipedia/commons/thumb/d/d6/Lock-gray-alt-2.svg/9px-Lock-gray-alt-2.svg.png")no-repeat;background-position:right .1em center}.mw-parser-output .cs1-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/thumb/a/aa/Lock-red-alt-2.svg/9px-Lock-red-alt-2.svg.png")no-repeat;background-position:right .1em center}.mw-parser-output .cs1-subscription,.mw-parser-output .cs1-registration{color:#555}.mw-parser-output .cs1-subscription span,.mw-parser-output .cs1-registration span{border-bottom:1px dotted;cursor:help}.mw-parser-output .cs1-hidden-error{display:none;font-size:100%}.mw-parser-output .cs1-visible-error{font-size:100%}.mw-parser-output .cs1-subscription,.mw-parser-output .cs1-registration,.mw-parser-output .cs1-format{font-size:95%}.mw-parser-output .cs1-kern-left,.mw-parser-output .cs1-kern-wl-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right,.mw-parser-output .cs1-kern-wl-right{padding-right:0.2em}


  2. ^ Hepler, P. K., S. M. Wick and S. M. Wolniak (1981). The structure and role of membranes in the mitotic apparatus. in: International Cell Biology 1980–1981, H.G. Schweiger, ed. Berlin: Springer-Verlag. pp. 673–686.CS1 maint: Multiple names: authors list (link)


  3. ^ Hepler, P. K.; B. A. Palevitz (1974). "Microtubules and microfilaments". Annual Review of Plant Physiology. 25: 309–362. doi:10.1146/annurev.pp.25.060174.001521.


  4. ^ abc Hepler, P. K.; B. A. Palevitz (1986). "Microtubules and microfilaments" (PDF). Current Contents (August 11, 1986) (32): 20. Retrieved October 7, 2016.


  5. ^ Hepler, J. R. (1922). Methods in Forcing Rhubarb: M.S. Thesis. University of Wisconsin. Retrieved October 7, 2016.


  6. ^ Hepler, Billy (2012). "America's Youngest Seed Grower" (PDF). Heritage Farm Companion (Summer): 6–9.


  7. ^ "A Bean Collector's Window". Retrieved October 18, 2016.


  8. ^ Hepler, Anna. "Anna Hepler Intricate Universe". Retrieved October 7, 2016.


  9. ^ "Membership Corner" (PDF) (31(5), 22). APBS News September/October 2004.


  10. ^ "Hepler Family (Pelham, MA)". UmassAmherst: MassWoods. Retrieved October 6, 2016.


  11. ^ "Kestrel Land Trust: Conserve the Valley You Love". Kestrel Land Trust. Retrieved October 6, 2016.


  12. ^ VandenBosch, K. A., W. Becker and B. A Palevitz (1996). "The natural history of a scholar and gentleman: A biography of Eldon H. Newcomb". Protoplasma. 195: 4–11. doi:10.1007/bf01279181. Retrieved October 7, 2016.CS1 maint: Multiple names: authors list (link)


  13. ^ ab Hepler, P. K., J. D. Pickett-Heaps and B. E. S. Gunning (2013). "Some retrospectives on early studies of plant microtubules". The Plant Journal. 75 (2): 189–201. doi:10.1111/tpj.12176. Retrieved October 7, 2016.CS1 maint: Multiple names: authors list (link)


  14. ^ Hepler, Peter K. (2016). "Founders' Review: The Cytoskeleton and Its Regulation by Calcium and Protons". Plant Physiology. 170: 3–22. doi:10.1104/pp.15.01506. PMC 4704593. Retrieved October 6, 2016.


  15. ^ "MBL Society Members". Marine Biological Laboratory. Retrieved October 6, 2016.


  16. ^ "Physiology 1981". History of the Marine Biological Laboratory. Retrieved October 6, 2016.


  17. ^ Hepler, P. K.; B. E. S. Gunning (1998). "Confocal fluorescence microscopy of plant cells". Protoplasma. 201 (3): 121–157. doi:10.1007/bf01287411. Retrieved October 6, 2016.


  18. ^ ab "AAAS Members Elected as Fellows". AAAS. Retrieved October 6, 2016.


  19. ^ ab "Members in the News". ASPB Newsletter 33(3), 26. April 2010. Retrieved October 6, 2016.


  20. ^ Hepler, P. K.; E. H. Newcomb (1967). "Fine structure of cell plate formation in the apical meristem of Phaseolus roots". Journal of Ultrastructure Research. 19 (5–6): 498–513. doi:10.1016/s0022-5320(67)80076-5. Retrieved October 7, 2016.


  21. ^ Palevitz, B. A.; P. K. Hepler (185). "Changes in dye coupling of stomatal cells of Allium and Commelina demonstrated by microinjection of Lucifer yellow". Planta. 164 (4): 473–479. doi:10.1007/bf00395962. Retrieved October 6, 2014.


  22. ^ Turgeon, R.; P. K. Hepler (1989). "Symplastic continuity between mesophyll and companion cells in minor veins of mature Cucurbita pepo L. leaves". Planta. 179 (1): 24–31. doi:10.1007/bf00395767. Retrieved October 6, 2016.


  23. ^ Zeiger, E.; P. K. Hepler (1976). "Production of Guard Cell Protoplasts from Onion and Tobacco". Plant Physiology. 58: 492–498. doi:10.1104/pp.58.4.492. PMC 543252. Retrieved October 6, 2016.


  24. ^ Zeiger, E., W. Moody, P. Hepler and F. Varela (1977). "Light-sensitive membrane potentials in onion guard cells". Nature. 270: 270–271. doi:10.1038/270270a0. Retrieved October 6, 2016.CS1 maint: Multiple names: authors list (link)


  25. ^ Zeiger, E.; P. K. Hepler (1977). "Light and stomatal function: blue light stimulates swelling of guard cell protoplasts". Science. 196 (4292): 887–889. doi:10.1126/science.196.4292.887. Retrieved October 6, 2016.


  26. ^ Zeiger, E.; P. K. Hepler (1979). "Blue light-induced, intrinsic vacuolar fluorescence in onion guard cells". Journal of Cell Science. 37: 1–10. Retrieved October 6, 2016.


  27. ^ Zhang, D., P. Wadsworth, and P. K. Hepler (1990). "Microtubule dynamics in living dividing cells: Confocal imaging of microinjected fluorescent brain tubulin". Proc. Natl. Acad. Sci. USA. 87: 8820–8824. doi:10.1073/pnas.87.22.8820. PMC 55051. PMID 11607116.CS1 maint: Multiple names: authors list (link)


  28. ^ Zhang, D., P. Wadsworth and P. K. Hepler (1993). "Dynamics of microfilaments are similar, but distinct from microtubules during cytokinesis in living, dividing plant cells". Cell Motility and the Cytoskeleton. 24: 151–155. doi:10.1002/cm.970240302. Retrieved October 7, 2016.CS1 maint: Multiple names: authors list (link)


  29. ^ Valster, A. H., E. S. Pierson, Valenta, P. K. Hepler and A. M. C. Emons (1997). "Probing the Plant Actin Cytoskeleton during Cytokinesis and Interphase by Profilin Microinjection". The Plant Cell. 9: 1815–1824. doi:10.1105/tpc.9.10.1815. PMC 157024. Retrieved October 7, 2016.CS1 maint: Multiple names: authors list (link)


  30. ^ Vos, J. W., A. H. Valster and P. K. Hepler (1988). "Methods for Studying Cell Division in Higher Plants". Methods in Cell Biology. 61: 413–437. doi:10.1016/S0091-679X(08)61992-5. Retrieved October 7, 2016.CS1 maint: Multiple names: authors list (link)


  31. ^ Hepler, P. K.; J. Hush (1996). "Behavior of Microtubules in Living Plant Cells". Plant Physiology. 112: 455–461. doi:10.1104/pp.112.2.455. PMC 157968. PMID 12226402.


  32. ^ ab Lancelle, S. A., D. A. Callaham and P. K. Hepler (1986). "A method for rapid freeze fixation of plant cells". Protoplasma. 131: 153–165. doi:10.1007/bf01285037. Retrieved October 6, 2016.CS1 maint: Multiple names: authors list (link)


  33. ^ "Poems and Quotations About the MicroWorld". Microscopy Society of America. Retrieved October 6, 2016.


  34. ^ ab Ledbetter, M. C.; K. R. Porter (1963). "A 'microtubule' in plant cell fine structure". Journal of Cell Biology. 19: 239–250. doi:10.1083/jcb.19.1.239. PMC 2106853.


  35. ^ Newcomb, E. H. (1996). "A career in science: Fulfillment of a dream". Protoplasma. 195 (1–4): 1–3. doi:10.1007/bf01279180. Retrieved October 7, 2016.


  36. ^ Sinnott, E. W.; R. Bloch (1945). "The cytoplasmic basis of intercellular patterns in vascular differentiation". American Journal of Botany. 32: 151–156. doi:10.2307/2437535.


  37. ^ Hepler, P. K.; E. H. Newcomb (1964). "The Fine Structure of Young Tracheary Xylem Elements Arising by Redifferentiation of Parenchyma in Wounded Coleus Stem". Journal of Experimental Botany. 14 (3): 496–503. doi:10.1093/jxb/14.3.496. Retrieved October 6, 2016.


  38. ^ Green, P. B. (1962). "Mechanism for plant cellular morphogenesis". Science. 138: 1404–1405. doi:10.1126/science.138.3548.1404.


  39. ^ Torrey, J. G., D. E. Fosket and P. K. Hepler (1971). "Xylem Formation: A Paradigm of Cytodifferentiation in Higher Plants: Plant cells divide and differentiate under the control of changing hormone levels. Xylem offers a model tissue for the study of these cellular events". American Scientist. 59 (3): 338–352. JSTOR 27829621.CS1 maint: Multiple names: authors list (link)


  40. ^ Wasteneys, G. O.; F. Brandizzi (2013). "A Glorious Half-Century of Microtubules". The Plant Journal. 75 (2): 185–188. doi:10.1111/tpj.12260. Retrieved October 6, 2016.


  41. ^ Palevitz, B. A.; P. K. Hepler (1976). "Cellulose microfibril orientation and cell shaping in developing guard cells of Allium: The role of microtubules and ion accumulation". Planta. 132: 71–93. doi:10.1007/BF00390333. PMID 24424910.


  42. ^ Lancelle, S. A., M. Cresti and P. K. Hepler (1987). "Ultrastructure of the cytoskeleton in freeze-substituted pollen tubes of Nicotiana alata". Protoplasma. 140: 141–150. doi:10.1007/bf01273723. Retrieved October 6, 2016.CS1 maint: Multiple names: authors list (link)


  43. ^ Inoué, S.; A. Bajer (1961). "Birefringence in endosperm mitosis". Chromosoma. 12: 48–63. doi:10.1007/bf00328913.


  44. ^ Hepler, P. K.; W. T. Jackson (1968). "Microtubules and early stages of cell plate formation in the endosperm of Haemanthus katherinae Baker". Journal of Cell Biology. 38: 437–446. doi:10.1083/jcb.38.2.437.


  45. ^ Hepler, P. K. (1976). "The blepharoplast of Marsilea: Its de novo formation and spindle association". Journal of Cell Science. 21: 361–390. Retrieved October 6, 2016.


  46. ^ Weisenberg, R. C. (1972). "Microtubule formation in vitro in solutions containing low calcium concentration". Science. 177: 1104–1105. doi:10.1126/science.177.4054.1104. Retrieved October 6, 2016.


  47. ^ Hepler, P. K. (1980). "Membranes in the mitotic apparatus of barley cells". Journal of Cell Biology. 86: 490–499. doi:10.1083/jcb.86.2.490. Retrieved October 6, 2016.


  48. ^ Wick, S. M.; P. K. Hepler (1980). "Localization of Ca++-containing antimonate precipitates during mitosis". Journal of Cell Biology: 500–513. doi:10.1083/jcb.86.2.500. Retrieved October 6, 2016.


  49. ^ Wolniak, S. M., P. K. Hepler, and W. T. Jackson (1980). "Detection of the membrane-calcium distribution during mitosis in Haemanthus endosperm with chlorotetracycline". Journal of Cell Biology. 87: 23–32. doi:10.1083/jcb.87.1.23. Retrieved October 6, 2016.CS1 maint: Multiple names: authors list (link)


  50. ^ Hepler, P. K.; D. A. Callaham (1987). "Free calcium increases during anaphase in stamen hair cells of Tradescantia". Journal of Cell Biology. 105: 2137–2143. doi:10.1083/jcb.105.5.2137. Retrieved October 7, 2016.


  51. ^ Hepler, P. K. (1989). "Calcium transients during mitosis: Observations in flux". Journal of Cell Biology. 109: 2567–2573. doi:10.1083/jcb.109.6.2567. Retrieved October 7, 2016.


  52. ^ Zhang, D. H. (1990). "Regulation of anaphase chromosome motion in Tradescantia stamen hair cells by calcium and related signaling agents". Journal of Cell Biology. 111: 171–182. doi:10.1083/jcb.111.1.171. Retrieved October 6, 2016.


  53. ^ Zhang, D. H., P. Wadsworth, and P. K. Hepler (1990). "Microtubule dynamics in living dividing plant cells: Confocal imaging of microinjected fluorescent brain tubulin". Proc. Natl. Acad. Sci. USA. 87 (22): 8820–8824. doi:10.1073/pnas.87.22.8820. PMC 55051. PMID 11607116. Retrieved October 6, 2016.CS1 maint: Multiple names: authors list (link)


  54. ^ Zhang, D. H., P. Wadsworth and P. K. Hepler (1992). "Modulation of anaphase spindle microtubule structure in stamen hair cells of Tradescantia by calcium and related agents". Journal of Cell Science. 102 (1): 79–89. Retrieved October 6, 2016.CS1 maint: Multiple names: authors list (link)


  55. ^ Palevitz, B. A., J. F. Ash, and P. K. Hepler (1974). "Actin in the green alga, Nitella". Proc. Natl. Acad. Sci. USA. 71 (2): 363–366. doi:10.1073/pnas.71.2.363. PMC 388005. PMID 4592689.CS1 maint: Multiple names: authors list (link)


  56. ^ Palevitz, B. A.; P. K. Hepler (1975). "Identification of actin in situ at the ectoplasm-endoplasm interface of Nitella. Microfilament-chloroplast association". Journal of Cell Biology. 65 (1): 29–38. doi:10.1083/jcb.65.1.29. Retrieved October 6, 2016.


  57. ^ Kersey, Y. M., P. K. Hepler, B. A. Palevitz, and N. K. Wessells (1976). "Polarity of actin filaments in Characean algae". Proc. Natl. Acad. Sci. USA. 73 (1): 165–167. doi:10.1073/pnas.73.1.165. PMC 335861. PMID 1061112.CS1 maint: Multiple names: authors list (link)


  58. ^ Hepler, P. K. (2005). "Historical Perspective Essay: Calcium: a central regulator of plant growth and development". Plant Cell. 17 (8): 2142–55. doi:10.1105/tpc.105.032508. Retrieved October 6, 2016.


  59. ^ Miller, D. D., D. A. Callaham, D. J. Gross and P. K. Hepler (1992). "Free Ca2+ gradient in growing pollen tubes of Lilium". Journal of Cell Science. 101: 7–12. Retrieved October 7, 2016.CS1 maint: Multiple names: authors list (link)


  60. ^ Wilsen, K. L.; P. K. Hepler (2007). "Sperm Delivery in Flowering Plants: The Control of Pollen Tube Growth". BioScience. 57 (10): 835–844. doi:10.1641/b571006. Retrieved October 20, 2016.


  61. ^ P. K. Hepler; J. G. Kunkel; C. M. Rounds; L. J. Winship (2012). "Calcium entry into pollen tubes". Trends in Plant Science. 17 (1): 32–38. doi:10.1016/j.tplants.2011.10.007. Retrieved October 7, 2016.


  62. ^ Wayne, R.; P. K. Hepler (1984). "The Role of Calcium Ions in Phytochrome-mediated germination of spores of Onoclea sensibilis L." Planta. 160: 12–20. doi:10.1007/bf00392460. Retrieved October 7, 2016.


  63. ^ Wayne, R.; P. K. Hepler (1985). "Red Light Stimulates and Increase in Intracellular Calcium in the Spores of Onoclea sensibilis" (PDF). Plant Physiology. 77: 8–11. doi:10.1104/pp.77.1.8. Retrieved October 7, 2016.


  64. ^ Saunders, M. J.; P. K. Hepler (1982). "Calcium ionophore A23187 stimulates cytokinin-like mitosis in Funaria". Science. 217: 943–945. doi:10.1126/science.217.4563.943.


  65. ^ Saunders, M. J.; P. K. Hepler (1981). "Localization of membrane-associated calcium following cytokinin treatment in Funaria using chlortetracycline". Planta. 152: 272–281. doi:10.1007/bf00385156. Retrieved October 6, 2016.


  66. ^ Conrad, P. A.; P. K. Hepler (1988). "The effect of 1,4-dihydropyridines on the initiation and development of gametophore buds in the moss Funaria". Plant Physiology. 86: 684–687. doi:10.1104/pp.86.3.684. Retrieved October 6, 2016.


  67. ^ "Jeanette Siron Pelton Award". Botanical Society of America. Retrieved October 8, 2016.


  68. ^ "Hepler named fellow of American Society of Plant Biologists". UmassAmherst News & Media Relations. Retrieved October 6, 2016.


  69. ^ "Peter K. Hepler". AAAS. Retrieved October 8, 2016.


  70. ^ "Hepler wins national award for plant discoveries". UmassAmherst News & Media Relations. Retrieved October 6, 2016.


  71. ^ "RMS Honorary Fellows". Royal Microscopical Society. Retrieved October 6, 2016.


  72. ^ "Hepler Named Honorary Fellow of Royal Microscopical Society". UmassAmherst News & Media Relations. Retrieved October 6, 2016.


  73. ^ "Peter K. Hepler Research Scholarship". UmassAmherst. Retrieved October 6, 2016.


  74. ^ title=Plant Biology Annual Symposium History|url=https://gpls.cns.umass.edu/pb/symposium/history









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