The Role of Bacteria in Chemical Signals of Elephant Musth: Proximate Causes and Biochemical Pathways

  • Thomas E. Goodwin
  • Innocent H. Harelimana
  • Laura J. MacDonald
  • Daniel B. Mark
  • Aline Umuhire Juru
  • Qin Yin
  • James A. Engman
  • Randall A. Kopper
  • Cheryl F. Lichti
  • Samuel G. Mackintosh
  • James D. Shoemaker
  • Mark V. Sutherland
  • Alan J. Tackett
  • Bruce A. Schulte


The chemical analysis of urine tells much about the physiological status of mammals, and often reveals compounds that function as chemical signals to conspecifics. Such is the case with mature male African (Loxodonta africana) and Asian (Elephas maximus) elephants in which there is odoriferous drainage from the temporal gland and dribbling of urine during musth, a periodic state in which serum androgens are elevated, food intake typically decreases, and aggressiveness between male conspecifics increases. We have employed solid phase dynamic extraction (SPDE)/GC-MS to identify a series of alkan-2-ones, alkan-2-ols, and a few simple aromatic compounds that increase in abundance in musth elephant urine. The primary focus of this report is on the alkan-2-ones and their corresponding alkan-2-ols, specifically: (1) the probable biosynthesis of these compounds via a secondary pathway for fatty acid metabolism, (2) the proximate cause for their increased abundance in musth urine, and (3) the role of bacteria in the increased abundance of these compounds exogenously in aged urine.


Asian Elephant African Elephant Scent Gland Secondary Pathway Stainless Steel Spatula 
These keywords were added by machine and not by the authors. This process is experimental and the keywords may be updated as the learning algorithm improves.



Elephant urine samples were supplied by Disney’s Animal Kingdom, Jacksonville Zoo, Miami Metro Zoo, and Riddle’s Elephant and Wildlife Sanctuary. We are most grateful to the dedicated staff members at these facilities for their indispensable help. Preliminary studies related to elephant albumin sequencing were assisted by Rick Edmundson (UAMS) and Dustin Freyaldenhoven (Hendrix College), as well as Leighton Satterfield (Hendrix College) who also carried out the Schiff stain for carbohydrates in the uromodulin. Shelly Bradley (Hendrix College), Larry O’Kane (Agilent), and Ingo Christ (Autosampler Guys) provided much-needed expertise to keep the instruments running. Peggy Morrison and Matthew Windsor were patient and persistent in tracking down many references. We appreciate the assistance of Henderson State University undergraduate students in the Engman research group (Mark Castleberry, Shannon Fiser, Jonathan Shields, Lauren Story, and Charlotte Wetzlar) who helped by sequencing two strains of bacteria from elephant urine. We are grateful for valuable advice from Kevin Theis (Michigan State University), and from Scott and Heidi Riddle (Riddle’s Elephant and Wildlife Sanctuary). We thank Hendrix College for financial support via a Distinguished Professor grant and the Julia Mobley Odyssey Professorship to T.E.G., as well as grants to students through the Odyssey Program. Additional funding in the early stages of this research was provided by the U.S. National Science Foundation (Award Nos. 02-17062, -17068, and -16862 to B.A.S., T.E.G. and the late L.E.L. Rasmussen, respectively). We would like to acknowledge the UAMS Proteomics Facility for mass spectrometric support. This work at UAMS was supported by National Institutes of Health grants P30GM103450, P20GM103429, and UL1TR000039.


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Copyright information

© Springer International Publishing Switzerland 2016

Authors and Affiliations

  • Thomas E. Goodwin
    • 1
  • Innocent H. Harelimana
    • 1
  • Laura J. MacDonald
    • 1
  • Daniel B. Mark
    • 1
  • Aline Umuhire Juru
    • 1
  • Qin Yin
    • 1
  • James A. Engman
    • 2
  • Randall A. Kopper
    • 1
  • Cheryl F. Lichti
    • 3
  • Samuel G. Mackintosh
    • 4
  • James D. Shoemaker
    • 5
  • Mark V. Sutherland
    • 6
  • Alan J. Tackett
    • 4
  • Bruce A. Schulte
    • 7
  1. 1.Department of ChemistryHendrix CollegeConwayUSA
  2. 2.Department of BiologyHenderson State UniversityArkadelphiaUSA
  3. 3.Department of Pharmacology and ToxicologyUniversity of Texas Medical Branch at GalvestonGalvestonUSA
  4. 4.Department of Biochemistry and Molecular BiologyUniversity of Arkansas for Medical Sciences (UAMS)Little RockUSA
  5. 5.E. A. Doisy Department of Biochemistry and Molecular BiologySaint Louis University School of MedicineSt. LouisUSA
  6. 6.Department of BiologyHendrix CollegeConwayUSA
  7. 7.Department of BiologyWestern Kentucky UniversityBowling GreenUSA

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