Joseph G. Kunkel
Born: Oceanside, New York, Aug. 17, 1942
74 Stony Hill Rd.
Biology Department,
Amherst MA 01002 University of Massachusetts
Amherst MA 01003
Phone:(413) 253-3391 (H).       (413) 545-0468 (Office)   (413)577-4426 (Lab)
Email: URL

Columbia College, New York, New York, Zoology A.B.'64
Case-Western Reserve U., Cleveland, Ohio, Biology Ph.D. '68   Dissertation:  Control of Cockroach Development

Society Memberships:
AAAS; SICB; Sigma Xi (Yale Chapter); MBL Corp.

Awards and Honors:
NSF Graduate Fellow, Biology, Case Western Reserve University, 1967-68; R.C.A. Scholar in Chemistry, Columbia College, 1961-62; Columbia University Scholar, Columbia College, 1960-64; New York State Regents Scholar, Columbia College, 1960-64; Bnai Brith Scholar, Columbia College, 1960

Positions and Professional Experience:

Professional and Competitive Grants Received:

Postdoctoral and Research Associates:

Graduate and Undergraduate Students:
  • Robert A. McCarthy, B.S. '73
  • Mary E. Johnson, M. S. '74
  • Donna M. Lawler, B.S. '74, Honors Thesis.
  • Gary Shepard, B.S. '76
  • Raymond C. Duhamel, Ph.D. '77
  • William T. Haggerty, M. S. '77
  • John R. Storella, M.S. '80
  • Colleen Finnegan, B.A. Hampshire College '81
  • Kerry Grolman, B.S. '81
  • Don M. Wojchowski, Ph. D. Zoology/MCB '84
  • Patricia Ann Estes, M. S. '85
  • William D. Shannon, M. S. '89
  • Myrta Oterro, B.S. '92
  • Sandra Torres, B.S. '92
  • Yujun Zhang, Ph. D. '92
  • Jeffrey Geller, B.S. '92 EE, Senior Honors Thesis
  • Raquell E. Dompenciel, Ph. D. MCB '93
  • Andrea Diss, Ph. D. Entomology '96
  • Anand R. Iyengar, Ph. D. MCB '96
  • Ruth Hartling, Ph. D. MCB '99
  • Ellen E. Faszewski, Ph. D. Biology '99
  • Brian R. Bettencourt, B.S.'96
  • Lan Tran B.S.'97
  • Ainex Baez B.S.'98, Honors Research
  • Paul Sonenblum, B.S.'99
  • Mike Pelak, B.S.'99
  • Edgar Rodriquez, B.S.'99
  • Janel Cho, B.S.'99
  • Brandon LaGrange, B.S. '99
  • Rahul Sharma, B.S. '99
  • Ray Moniz, B.S. '00
  • Jenny Mirabal, B.S. '00
  • Jennifer Schnorbus, '01
  • Winnie Pong '02, Honors Student
  • Jeff Xu



Hagedorn, HH and JG Kunkel. 1979. Vitellogenin & Vitellin in Insects. Annu Rev Entomol 24:475-505.

Kunkel JG. 1981. A minimal model of metamorphosis: fat body competence to respond to Juvenile Hormone. in Current Trends in INSECT ENDOCRINOLOGY AND NUTRITION, eds. G Bhaskaran, M Friedman and J Rodrigues, Plenum Press, New York, pp107-130.

Kunkel JG. 1981. Cockroach Regeneration. Chapter 16 in THE AMERICAN COCKROACH, eds. WJ Bell and KG Adiyodi, Chapman and Hall, London, pp 427-443.

Kunkel JG and JH Nordin. 1985. Yolk Proteins. in COMPREHENSIVE INSECT PHYSIOLOGY, BIOCHEMISTRY AND PHARMACOLOGY. Chapter 4, Vol.I, eds. GA Kerkut and LI Gilbert, Pergamon Press, pp 83-111.

Kunkel JG.  1988.  Analytical Immunological Techniques. Chapter I in IMMUNOLOGICAL TECHNIQUES: ARTHROPODS. Edited by LI Gilbert. Springer Verlag, pp 1-41.

Leonard DE and JG Kunkel. 1989. Nutritional Ecology: Lymantria dispar as a model system for study of serum storage proteins. In LYMANTRIDAE: A COMPARISON OF FEATURES OF NEW AND OLD WORLD TUSSOCK MOTHS, PROCEEDINGS. USDA Gen. Tech. Report NE-123: 367-80.

Kunkel JG. 1990. Review of OOCYTE GROWTH AND MATURATION by T. A. Dettlaff & S. G. Vassetzky. Quart Rev Biol Sci 65:506.

Telfer WH and JG Kunkel. 1991.  The function and evolution of insect storage hexamers. Annu Rev Entomol 36: 205-28.

Kunkel, JG, L-Y Lin, Y Xu, AMM Prado, JA Feijó, PP Hwang and PK Hepler.  2001.  The strategic use of Good buffers to measure proton gradients about growing pollen tubes. In CELL BIOLOGY OF PLANT AND FUNGAL TIP GROWTH. Edited by A Geitman.  Amsterdam: IOS Press, pp. 14pp. (in press).


Kunkel JG 1966. Development and the availability of food in the German Cockroach, Blattella germanica (L.) J.Insect Physiol. 12:227-235.

Kunkel JG 1973. Gonadotrophic effect of Juvenile Hormone in Blattella germanica. A rapid, simple quantitative bioassay. J.Insect Physiol. 19: 1285-1297.

Kunkel JG and DM Lawler. 1974. Larval-specific serum protein in the order Dictyoptera I. Immunological characterization in larval Blattella germanica and cross-reaction throughout the order. Comp.Biochem.Physiol. 47B: 697-710.

Kunkel JG 1975a. Larval-specific serum protein in the order Dictyoptera - II. Antagonistic effects of ecdysone and regenerationon LSP concentration in the hemolymph of the Oriental Cockroach, Blatta orientalis. Comp.Biochem.Physiol. 51B: 177-188.

Kunkel JG 1975b. Cockroach Molting I. Temporal organization of events during the molting cycle of Blattella germanica. Biol.Bull. 148: 259-273.

Kunkel JG and ML Pan. 1976. Selectivity of yolk protein uptake: Comparison of vitellogenins of two insects. J.Insect Physiol. 22: 809-818.

Kunkel JG 1977. Cockroach Molting II. The nature of regeneration-induced delay of molting hormone secretion. Biol.Bull.153: 145-162.

Duhamel RC and JG Kunkel. 1978. A molting rhythm for serum proteins of the cockroach, Blatta orientalis. Comp.Biochem.Physiol.60B: 333-337.

Greenberg SLW, JG Kunkel and AM Stuart. 1978. Vitellogenesis in a primitive termite, Zootermopsis angusticollis (Hagen). Biol.Bull. 155: 336-346.

Hagedorn, HH, JG Kunkel and G Wheelock. 1978. The specificity of an antiserum against mosquito vitellogenin and its use in a radio-immunological precipitation assay for protein synthesis. J.Insect Physiol. 24: 481-489.

Greenberg SLW. and AM Stuart. 1979. The influence of group size on ovarian development in adult and neotenic reproductives of the termite Zootermopsis angusticollis (Hagen)(Hodotermitidae). Int. J. Invert. Rep. 1: 99-108. (student paper)

Cherry LM, SM Case, JG Kunkel and AC Wilson. 1979. Comparisons of frogs, humans and chimpanzees. Science 204: 435.

Kunkel JG, LM Cherry, SM Case and AC Wilson. 1980. M-Statistics and morphometric divergence. Science 208: 1060-1061.

Kunkel JG, G Shepard, RA McCarthy, DB Ethier and JH Nordin. 1981. Concanavalin A reactivity and carbohydrate structure of Blattella germanica vitellin. Insect Biochem. 10: 703-714.

Cherry LM, SM Case, JG Kunkel, JS Wyles and AC Wilson. 1982. Body shape metrics and organismal evolution. Evolution 36: 914-933.

Duhamel RC and JG Kunkel. 1983. Cockroach larval-specific protein (LSP), a tyrosine-rich serum protein. J.Biol.Chem. 258: 14461-14465.

Wyles JS, JG Kunkel and AC Wilson. 1983. Birds, behavior, and anatomical evolution. Proc.Natl.Acad.Sci.USA 80: 4394-4397.

Wilson AC, JG Kunkel and JS Wyles. 1984. Morphological distance: An encounter between two perspectives in evolutionary biology. Evolution 38: 1156-1159.

Nordin JH, C Gochoco, D Wojchowski and JG Kunkel. 1984. Limited and heterogenous processing of high mannose oligosaccharides of insect vitellins. Comp.Biochem.Physiol. 79B: 379-390.

Storella JR, DM Wojchowski & JG Kunkel. 1985. Structure and embryonic degradation of two native vitellins in the cockroach, Periplaneta americana. Insect Biochem. 15: 259-275.

Kunkel JG, R Koenig, H Kindle and B Lanzrein. 1986. The role of ions in vitellogenesis and patterning in insect oocytes. ADVANCES IN INVERTEBRATE REPRODUCTION 4: 101-8.

Wojchowski DM, PA Parsons, JH Nordin and JG Kunkel. 1986. Processing of Provitellogenin in Insect Fat Body: a role for High-Mannose Oligosaccharide. Developmental Biol. 116: 422-430.

Kunkel JG. 1986. Dorsoventral currents are associated with vitellogenesis in cockroach ovarioles. in IONIC CURRENTS IN DEVELOPMENT ed. R Nuccitelli, Alan R Liss Publ., pp 165-72.

Wojchowski DM and JG Kunkel. 1987. Purification of two distinct oocyte vitellins and identification of their corresponding vitellogenins in fat body and hemolymph of Blaberus discoidalis. Insect Biochem. 17: 189-198.

Duhamel RC and JG Kunkel. 1987. Molting cycle regulation of hemolymph protein clearance in cockroaches: Possible size-dependent mechanism. J.Insect Physiol. 33: 155-158.

Koenig R, JH Nordin, CH Gochoco and JG Kunkel. 1988 Studies on ligand recognition by vitellogenin receptors in follicle membrane preparations of the German cockroach Blattella germanica. Insect Biochemistry 18, 395-404.

Purcell JP, JG Kunkel and JH Nordin. 1988. Yolk hydrolase activities associated with polypeptide and oligosaccharide processing of Blattella germanica vitellin. Arch.Insect Bioch.Physiol. 9: 39-58.

Gochoco CH, Kunkel JG and Nordin JH. 1988. The high mannose oligosaccharides of Blattella germanica vitellin participate in its endocytosis in vivo and maintenance of its multimeric structure in vitro. Arch.Insect Biochem.Physiol.

Gochoco CH, JG Kunkel and JH Nordin. 1988. Experimental modifications of an insect vitellin affect its structure and uptake by oocytes. Arch.Insect Bioch.Physiol. 8: 179-199.

Koenig R, H Kindle, JG Kunkel and B Lanzrein. 1988. Vitellogenisis in the cockroach Nauphoeta cinerea: Separation of two classes of ovarian binding sites and calcium effects on binding and uptake. Arch.Insect Biochem.Physiol. 9: 323-337.

Purcell JP, TM Quinn, JG Kunkel and JH Nordin. 1989. Correlation of yolk phosphatase expression with the programmed proteolysis of vitellin in Blattella germanica during embryonic development. Arch.Insect Bioch. Physiol. 9: 237-250.

Kunkel JG and E Bowdan. 1989. Modeling currents about vitellogenic oocytes of the cockroach, Blattella germanica. Biol. Bull.176(S): 96-102.

Kunkel JG, E Bowdan, H Kindle and B Lanzrein. (1989). An inhibitory role of glucose in the vitellogenic process. In REGULATION OF INSECT REPRODUCTION IV Ed. by M Tonner, T Soldan and B Bennettova. Academia Publishing House of the Czechoslovak Academy of Sciences, pp 85-86.

Bowdan E & JG Kunkel. Patterns of ionic currents around the developing oocyte of the German Cockroach, Blattella germanica. Developmental Biology 137: 266-275. Cover photo.

Kunkel JG, C Grossniklaus-Buergin, ST Karpells and B Lanzrein. Arylphorin of Trichoplusia ni: Characterization and parasite induced precocious increase in titer. Arch.Insect Bioch. Physiol. (in press).

Karpells SA, DE Leonard and JG Kunkel. Cyclic fluctuations in arylphorin, the principal serum storage protein in the gypsy moth, Lymantria dispar. Insect Biochemistry 20:73-82.

Anderson M & JG Kunkel. 1990. Cleaning insect oocytes by dissection and enzyme treatment. Tiss. Cell 22:349-358.

Sigel E, R Baur, JG Kunkel, H Kindle and B Lanzrein. 1990. Demonstration of a voltage dependent calcium current in follicles of the cockroach Nauphoeta cinerea. Invert Reprod Dev 18:159-164.

Zhang Y, and JG Kunkel. 1991. High abundance calmodulin from Blattella germanica eggs binds to vitellin subunits but disappears during vitellin utilization. Insect Biochem. 22:293-304.

Dompenciel RE, CM Yin, DE Leonard, JG Kunkel. (1992) Larval allatectomy affects vitellogenin accumulation in oocytes along ovarioles of the gypsy moth, Lymantria dispar. In: Regulation of Insect Reproduction V. (eds. B Bennetova, I Gelbic, T Soldan)  Academia Publishing Czechoslovak Academy of Science; Prague, CZ.

Zhang Y. 1992. Characterization of insect calmodulin during oogenesis and embryogenesis of Blattella germanica. Dissertation, University of Massachusetts, Amherst, MA, 137.

Zhang Y, and JG Kunkel. 1992. Program of F-actin in the follicular epithelium during oogenesis of the German cockroach, Blattella germanica. Tiss. Cell 24:905-917.

Zhang Y, and JG Kunkel. 1994. Most egg calmodulin is a follicle cell contribution to the cytoplasm of the Blattella germanica oocyte. Dev. Biol. 160:513-521.

Keefe D, J Pepperell, P Rinaudo, JG Kunkel and PJS Smith. 1995.  Identification of calcium flux in single preimplantation mouse embryos with the calcium-sensitive vibrating probe. Biol Bull, 189, 200.

Iyengar AR, and JG Kunkel. 1995. Follicle cell calmodulin in Blattella germanica: Transcript accumulation during vitellogenesis is regulated by juvenile hormone. Dev. Biol. 170:314-320.

Hartling RC, JJ Pereira, JG Kunkel. 1997. Characterization of a heat-stable fraction of lipovitellin and development of an immunoassay for vitellogenin and yolk protein in winter flounder (Pleuronectes americanus). J. Exp. Zool. 278:156-166

Feijo JA, Sainhas J, Hackett GR, Kunkel JG, Hepler PK. 1999. Growing pollen tubes possess a constitutive alkaline band in the clear zone and a growth-dependent acidic tip. J. Cell Biol. 144:483-96

Cardenas L, Feijo JA, Kunkel JG, Sanchez F, Holdaway-Clarke T, Hepler PK, Quinto C. 1999. Rhizobium nod factors induce increases in intracellular free calcium and extracellular calcium influxes in bean root hairs. Plant J 19:347-52

Hartling RC and Kunkel JG. 1999. Developmental fate of the yolk protein lipovitellin in embryos and larvae of winter flounder, Pleuronectes americanus. J Exp Zool 284:686-95.

Roy SJ, TL Holdaway-Clarke, GR Hackett, JG Kunkel, EM Lord, PK Hepler. 1999.  Uncoupling secretion and tip growth in lily pollen tubes: evidence for the role of calcium in exocytosis. Plant J, 19:379-86.

Cardenas L, TL Holdaway-Clarke, F Sanchez, C Quinto, JA Feijo, JG Kunkel, PK Hepler.  2000.  Ion changes in legume root hairs responding to Nod factors. Plant Physiol, 123:443-52.

Feijo JA, J Sainhas, T Holdaway-Clarke, MS Cordeiro, JG Kunkel and PK Hepler.  2001. Cellular oscillations and the regulation of growth: the pollen tube paradigm. Bioessays 23: 86-94.


Kunkel JG, DB Ethier and JH Nordin. 1978. Carbohydrate structure and immunochemical properties of Blattella vitellin. Fed. Proc. 37: 948.

Kunkel JG. 1979. Stages of vitellogenic competence in the cockroach. American Zool. 19:999.

Storella JR and JG Kunkel. 1979. Processing and utilization of vitellin in the cockroach egg. American Zool. 19: 999.

Cherry LM, JG Kunkel, SM Case, JS Wyles and AC Wilson. 1980. The trouble with Mahalanobis Distance. 14th International Numerical Taxonomy Meetings, Norman, Oklahoma. *

Kunkel JG. 1980. Metamorphosis of cockroach fat body to a vitellogenicly competent state. 3rd International Vitellogenin Conference, Kandersteg, Switzerland. *

Kunkel JG. 1980. Size and shape predictions of cockroach reproductive metamorphosis. American Zool. 20: 937.

Storella JR and JG Kunkel. 1981. Two vitellogenins in the cockroach, Periplaneta americana. Cell Biology 91: 23016.

Parsons PA, K Grolman and JG Kunkel. 1981. Cell fractionation of low urate fat body from cockroaches. Cell Biology 91: 1057.

Nordin JH and JG Kunkel. 1982. Size heterogeneity of the high mannose oligosaccharide of insect vitellin. Fed.Proc. 41: 886.

Kunkel JG and DM Wojchowski. 1982. Development of Vitellogenic competence in insects: a sexually dimorphic event. Proceedings of The Ninth International Symposium on Comparative Endocrinology. Hong Kong, December 1981. Hong Kong University Press., p13.

Kunkel JG 1982. Metamorphosis of the cockroach fat body- A biochemically tractable problem. Eastern Branch Meetings of the Entomological Society of America.

Wojchowski DM and JG Kunkel. 1982. Juvenile Hormone receptors during metamorphosis in the cockroach. Tenth Annual New England Endocrinology Conference. Mount Holyoke, Mass. *

Wojchowski DM and JG Kunkel. 1983. Developmental heterogeneity of Juvenile Hormone binding sites in insect fat body cytosol. Fed.Proc. 42: 709.

Kunkel JG, DM Wojchowski, CH Gochoco and JH Nordin. 1983. Tunicamycin inhibition of high mannose oligosaccharide attachment to an insect vitellogenin abolishes its normal processing and secretion from the fat body. Fed.Proc. 42: 1904.

Kunkel JG, PA Estes, CH Gochoco, JP Purcell, DM Wojchowski and JH Nordin. 1984. Vitellogenin of the German Cockroach, Blattella germanica: Structure and function of high mannose oligosaccharide in secretion, uptake and storage. Proceedings of the XVII International Congress of Entomology. Hamburg, FRG.

Gochoco CH, JG Kunkel and JH Nordin. (1984). The role of oligosaccharide in the endocytosis of an insect vitellin by the oocyte. Meetings of the American Society for Cell Biology, Kansas City, MO.

Nordin JH, CH Gochoco, JP Purcell, DM Wojchowski and JG Kunkel. 1984. Evidence that the high mannose oligosaccharide of an insect glycoprotein functions in the maintenance of its multimeric structure in vitro and assembly and secretion in vivo. Twelfth International Carbohydrate Symposium, Utrecht, The Netherlands.

Kunkel JG, R Koenig and B Lanzrein. 1986. The role of ions in vitellogenesis and patterning in insect oocytes. IV International Symposium on Invertebrate Reproduction, Lille, France.

Koenig R, JH Nordin and JG Kunkel. 1986. The role of the carbohydrate component in binding of vitellin to oocyte membranes of the German Cockroach, Blattella germanica. IV International Symposium on Invertebrate Reproduction, Lille, France.

Gochoco CH, JG Kunkel & JH Nordin. 1986. Development and partial characterization of an in vitro system for studying endocytosis of insect vitellogenin by oocytes. Fed.Proc. 45: 1911.

Karpells ST, JG Kunkel and DE Leonard. 1986. Arylphorin of the gypsy moth, Lymantria dispar: Purification properties and developmental profile. Entomological Society of America, Reno, Nevada, A1068.

Purcell J, JG Kunkel and JH Nordin. 1986. Evidence that a yolk phosphatase functions in developmentally-linked proteolysis of Blattella germanica vitellin. Entomological Society of America, Reno, Nevada, DP24.

Koenig R, JH Nordin, JG Kunkel and B Lanzrein. 1987. Demonstration of two classes of vitellogenin binding sites. in Abst. AAAS meeting.

Koenig R, H Kindle, JG Kunkel and B Lanzrein. 1987. The effect of calcium on receptor-mediated endocytosis of vitellogenin in cockroach follicles in-vitro. Union of Swiss Societies of Experimental Biology, Lausanne, Switzerland. Experientia 43: 688. (Abstract)

Kunkel JG. (1988). Evolution of composition and function of insect serum storage proteins. Proc.XVIII Int.Cong.Ent., Vancouver BC, p73 (Abs).

Karpells ST, RE Dompencial-Rivera, DE Leonard and JG Kunkel. (1988). Arylphorin of the gypsy moth, Lymantria dispar: Two distict roles in development. Proc.XVIII Int.Cong.Ent., Vancouver BC, p115 (Abs).

Kunkel JG. (1988). Role of ions in the development of pattern in early development. Program Abstacts & History CWRU Dept. Biol. Centennial, p22. *

Zhang Y and JG Kunkel (1989). Calcium binding proteins in an insect oocyte. Biol. Bull. 176: 71 (abstract), best graduate student poster award.

Bowdan E and JG Kunkel (1989). Patterns of ionic currents about developing cockroach oocyte. Biol.Bull. 176:68 (abstract).

Dompenciel-Rivera RE, Kunkel JG, Leonard DE. (1990) Diversification of yolk proteins: An insect embryonic strategy? In: Molecular Insect Science (eds. HH Hagedorn, JG Hildebrand, MG Kidwell, JH Law), pp 299. Plenum, NY.

Hartling RC, Kunkel JG. (1995) Proteolytic cleavage of yolk protein during flounder (Pleuronectes americanus) development: characterization of lipovitellin from eggs and embryos. Mol Biol Cell, 6, 321a.

Hartling RC, Pereira JJ, Kunkel JG. (1995) Processing and degradation of lipovitellin during embryonic and larval development of the winter flounder (Pleuronectes americanus). FASEB J, 9, A81.

Holdaway-Clarke TL, Hackett GA, Kunkel JG, Hepler PK, Feijo JA. (1998) Oscillations of cell expansion rate, cytoplasmic calcium, and calcium influx in the pollen tube. JOURNAL OF GENERAL PHYSIOLOGY, 112: 94.

Kunkel JG, J Bohannon, R Sharma, and J Zydlewski.  (1999) Serum and slime vitellogenin in the Atlantic cod, Gadus morhua, and its relation to ovarian development. Am. Zool. 39:A119.

Zydlewski J, McCormick SD, Kunkel JG. (1999) Early seawater entry of juvenile American shad may be advantageous. Am. Zool. 39:A546.

Zydlewski J, McCormick SD, Kunkel JG. (2000) The Ecological Physiology of Juvenile American shad (Alosa sapidissima). In: First conference on European shads, Centre Condorcet of Pessac, Bordeaux, France.