Publish with BioMed Central and every
scientist can read your work free of charge
"BioMed Central will be the most significant development for
disseminating the results of biomedical research in our lifetime."
Sir Paul Nurse, Cancer Research UK
Your research papers will be:
available free of charge to the entire biomedical community
peer reviewed and published immediately upon acceptance
cited in PubMed and archived on PubMed Central
yours — you keep the copyright
Submit your manuscript here:
BMC Biology 2009, 7:14 http://www.biomedcentral.com/1741-7007/7/14
Page 14 of 14
(page number not for citation purposes)
ses were done using MultiExperiment Viewer http://
www.tm4.org/mev.html. Gene annotation was done
using the previously published ESTIMA monarch EST
The authors declare that they have no competing interests.
All authors contributed to experimental design, execution,
data analysis and writing the paper. All authors have read
and approved the final manuscript.
We thank Lauren Foley and Danielle Metterville for technical assistance;
Carol Cullar and Fred Gagnon for supplying butterflies; and Adriana Briscoe
and Christine Merlin for comments and discussions. Supported in part by
NIH grant R01GM086794 and NSF grant IOB-0646060.
1. Brower LP: Understanding and misunderstanding the migra-
tion of the monarch butterfly (Nymphalidae) in North
America: 1857–1995. J Lepid Soc 1995, 49:304-385.
2. Herman WS: Endocrine regulation of post eclosion enlarge-
ment of the male and female reproductive glands in mon-
arch butterflies. Gen Comp Endocrinol 1975, 26:534-540.
3. Herman WS, Tatar M: Juvenile hormone regulation of longevity
in the migratory monarch butterfly. Proc Biol Soc 2001,
4. Froy O, Gotter AL, Casselman AL, Reppert SM: Illuminating the
circadian clock in monarch butterfly migration. Science 2003,
5. Mouritsen H, Frost BJ: Virtual migration in tethered flying mon-
arch butterflies reveals their orientation mechanisms. Proc
Natl Acad Sci USA 2002, 99:10162-10166.
6. Perez SM, Taylor OR, Jander R: A sun compass in monarch but-
terflies. Nature 1997, 387:29.
7. Brower L: Monarch butterfly orientation: missing pieces of a
magnificent puzzle. J Exp Biol 1996, 199:93-103.
8. Herman WS, Brower LP, Calvert WH: Reproductive tract devel-
opment in monarch butterflies overwintering in California
USA and Mexico. J Lepid Soci 1989, 43:50-58.
9. Kanz JE: The orientation of migrant and nonmigrant monarch
butterflies danaus-plexippus. Psyche (Cambridge) 1977,
10. Perez SM, Taylor OR: Monarch butterflies' migratory behavior
persists despite changes in environmental conditions. In Mon-
arch Butterfly Biology & Conservation Edited by: Oberhauser KS, Solen-
sky MJ. Ithaca: Cornell University Press; 2004:85-88.
11. Zhu H, Casselman A, Reppert SM: Chasing migration genes: a
brain expressed sequence tag resource for summer and
migratory monarch butterflies (Danaus plexippus). PLoS ONE
12. Robinson GE: Effects of a juvenile hormone analog on honey
bee apis-mellifera-ligustica foraging behavior and alarm phe-
romone production. J Insect Physiol 1985, 31:277-282.
13. Stanewsky R: Genetic analysis of the circadian system in Dro-
sophila melanogaster and mammals. J Neurobiol 2003,
14. Shinoda T, Itoyama K: Juvenile hormone acid methyltrans-
ferase: a key regulatory enzyme for insect metamorphosis.
Proc Natl Acad Sci USA 2003, 100:11986-11991.
15. Li S, Ouyang YC, Ostrowski E, Borst DW: Allatotropin regulation
of juvenile hormone synthesis by the corpora allata from the
lubber grasshopper, Romalea microptera. Peptides 2005,
16. Sarov-Blat L, So WV, Liu L, Rosbash M: The Drosophila takeout
gene is a novel molecular link between circadian rhythms
and feeding behavior. Cell 2000, 101:647-656.
17. Barker JF, Herman WS: Effect of photoperiod and temperature
on reproduction of the monarch butterfly danaus-plexippus.
J Insect Physiol 1976, 22:1565-1568.
18. Goehring L, Oberhauser KS: Effects of photoperiod, tempera-
ture, and host plant age on induction of reproductive dia-
pause and development time in Danaus plexippus. Ecol
Entomol 2002, 27:674-685.
19. Borland J, Johnson CC, Crumpton TW III, Thomas M, Altizer SM,
Oberhauser KS: Characteristics of fall migratory monarch but-
terflies, Danaus plexippus, in Minnesota and Texas. In Mon-
arch Butterfly Biology & Conservation Edited by: Oberhauser KS,
Solensky MJ. Ithaca: Cornell University Press; 2004.
20. Prysby MD, Oberhauser KS: Temporal and geographic variation
in monarch densities: citizen scientists document monarch
population patterns. In Monarch Butterfly Biology & Conservation
Edited by: Oberhauser KS, Solensky MJ. Ithaca: Cornell University
21. Whitfield CW, Cziko AM, Robinson GE: Gene expression profiles
in the brain predict behavior in individual honey bees. Science
22. Reppert SM, Zhu H, White RH: Polarized light helps monarch
butterflies navigate. Curr Biol
23. Batschelet E: Circular statistics in biology. London, New York:
Academic Press; 1981.
24. Nuwaysir EF, Huang W, Albert TJ, Singh J, Nuwaysir K, Pitas A, Rich-
mond T, Gorski T, Berg JP, Ballin J, McCormick M, Norton J, Pollock
T, Sumwalt T, Butcher L, Porter D, Molla M, Hall C, Blattner F, Suss-
man MR, Wallace RL, Cerrina F, Green RD: Gene expression anal-
ysis using oligonucleotide arrays produced by maskless
photolithography. Genome Res 2002, 12:1749-1755.
Additional file 1
Sex-specific orientation genes.
Click here for file
Additional file 2
Sex-specific juvenile hormone-regulated genes.
Click here for file