Nancy A. Moran


Moran lab publications

Nancy Moran's Google Scholar page

engineered bee gut bacteria


Villabona N, Moran N, Hammer T, Reyes A. 2023. Conserved, yet disruption-prone, gut microbiomes in neotropical bumblebees. mSphere  e0013923. doi: 10.1128/msphere.00139-23. [Online ahead of print.]

Huang Q, Lariviere PJ, Powell JE, Moran NA. 2023. Engineered gut symbiont inhibits microsporidian parasite and improves honey bee survival. Proc Natl Acad Sci U S A. 120(25):e2220922120. doi: 10.1073/pnas.2220922120.

Elston KM, Phillips LE, Leonard SP, Young E, Holley JC, Ahsanullah T, McReynolds B, Moran NA, Barrick JE. 2023.  The Pathfinder plasmid toolkit for genetically engineering newly isolated bacteria enables the study of Drosophila-colonizing Orbaceae. ISME Commun . 3(1):49. doi: 10.1038/s43705-023-00255-3.

Motta EVS, Arnott RLW, Moran NA. 2023. Caffeine consumption helps honey bees fight a bacterial pathogen. Microbiol Spectr . 11(3):e0052023. doi: 10.1128/spectrum.00520-23.

Powell JE, Lau P, Rangel J, Arnott R, De Jong T, Moran NA. 2023. The microbiome and gene expression of honey bee workers are affected by a diet containing pollen substitutes. PLoS One . 18(5):e0286070. doi: 10.1371/journal.pone.0286070

Motta EVS, Moran NA. 2023. The effects of glyphosate, pure or in herbicide formulation, on bumble bees and their gut microbial communities. Sci Total Environ. 872:162102. doi: 10.1016/j.scitotenv.2023.162102.

Hammer TJ, Easton-Calabria A, Moran NA. 2023. Microbiome assembly and maintenance across the lifespan of bumble bee workers. Mol Ecol . 32(3):724-740. doi: 10.1111/mec.16769.


Motta EVS, Gage A, Smith TE, Blake KJ, Kwong WK, Riddington IM, Moran NA. 2022. Host-microbiome metabolism of a plant toxin in bees. eLife11:e82595. doi: 10.7554/eLife.82595

Li Z, Li Y, Xue AZ, Dang V, Renee Holmes V, Spencer Johnston J, Barrick JE, Moran NA. 2022. The genomic basis of evolutionary novelties in a leafhopper. Mol Biol Evol . 39(9):msac184. doi: 10.1093/molbev/msac184.

Holley JC, Jackson MN, Pham AT, Hatcher SC, Moran NA. 2022. Carpenter bees (Xylocopa) harbor a distinctive gut microbiome related to that of honey bees and bumble bees. Appl Environ Microbiol. 88(13):e0020322. doi: 10.1128/aem.00203-22.

Smith TE, Li Y, Perreau J, Moran NA. 2022. Elucidation of host and symbiont contributions to peptidoglycan metabolism based on comparative genomics of eight aphid subfamilies and their Buchnera. PLoS Genet. 18(5):e1010195. doi: 10.1371/journal.pgen.1010195.

Motta EVS, Powell JE, Leonard SP, Moran NA. 2022. Prospects for probiotics in social bees. Philos Trans R Soc Lond B Biol Sci . 377(1853): 20210156. doi: 10.1098/rstb.2021.0156.

Li Y, Leonard SP, Powell JE, Moran NA. 2022. Species divergence in gut-restricted bacteria of social bees. Proc Natl Acad Sci U S A . 2022 119(18):e2115013119. doi: 10.1073/pnas.2115013119.

Nakabachi A, Moran NA. 2022. Extreme polyploidy of Carsonella, an organelle-like bacterium with a drastically reduced genome. Microbiol Spectr. 10(3):e0035022. doi: 10.1128/spectrum.00350-22

Busby TJ, Miller CR, Moran NA, Van Leuven JT. 2022. Global Composition of the Bacteriophage Community in Honey Bees. mSystems 26;7(2):e0119521. doi: 10.1128/msystems.01195-21

Motta EVS, Powell JE, Moran NA. 2022. Glyphosate induces immune dysregulation in honey bee. Anim Microbiome 4(1):16. doi: 10.1186/s42523-022-00165-0

Blaxter M, Archibald JM, Childers AK, Coddington JA, Crandall KA, Di Palma F, Durbin R, Edwards SV, Graves JAM, Hackett KJ, Hall N, Jarvis ED, Johnson RN, Karlsson EK, Kress WJ, Kuraku S, Lawniczak MKN, Lindblad-Toh K, Lopez JV, Moran NA, Robinson GE, Ryder OA, Shapiro B, Soltis PS, Warnow T, Zhang G, Lewin HA. 2022. Why sequence all eukaryotes? Proc Natl Acad Sci U S A. 119(4):e2115636118. doi: 10.1073/pnas.2115636118

Perreau J, Moran NA. 2022. Genetic innovations in animal-microbe symbioses. Nat Rev Genet. 23(1): 23-39. doi: 10.1038/s41576-021-00395-z.


Moran NA. 2021. Microbe profile: Buchnera aphidicola: ancient aphid accomplice and endosymbiont exemplar. Microbiology (Reading). 167(12). doi: 10.1099/mic.0.001127

Smith TE, Lee M, Person MD, Hesek D, Mobashery S, Moran NA. 2021. Horizontal-acquisition of a promiscuous peptidoglycan-recycling enzyme enables aphids to influence symbiont cell wall metabolism. mBio. 12(6):e0263621. doi: 10.1128/mBio.02636-21

Hammer TJ, Moran NA. 2021. The gut microbiota of bumblebees. Insectes Soc. 68 (2-3). doi: 10.1007/s00040-021-00837-1

Steele MI, Motta EVS, Gattu T, Martinez D, Moran NA. 2021. The gut microbiota protects bees from invasion by a bacterial pathogen. Microbiol Spectr. 9(2):e0039421. doi: 10.1128/Spectrum.00394-21

Perreau J, Zhang B, Maeda GP, Kirkpatrick M, Moran NA. 2021. Strong within-host selection in a maternally inherited obligate symbiont: Buchnera and aphids. Proc Natl Acad Sci U S A. 118(35):e2102467118. doi: 10.1073/pnas.2102467118

Powell JE, Carver Z, Leonard SP, Moran NA. 2021. Field-realistic tylosin exposure impacts honey bee microbiota and pathogen susceptibility, which is ameliorated by native gut probiotics. Microbiol Spectr. 9(1):e0010321. doi: 10.1128/Spectrum.00103-21.

Cerqueira AES, Hammer TJ, Moran NA, Santana WC, Kasuya MCM, da Silva CC. 2021. Extinction of anciently associated gut bacterial symbionts in a clade of stingless bees. ISME J. 15(9):2813-2816. doi: 10.1038/s41396-021-01000-1.

Steele MI, Moran NA. 2021. Evolution of interbacterial antagonism in bee gut microbiota reflects host and symbiont diversification. mSystems 6(3):e00063-21. doi: 10.1128/mSystems.00063-21

Schepers MJ, Yelland JN, Moran NA, Taylor DW. 2021. Isolation of the Buchnera aphidicola flagellum basal body complexes from the Buchnera membrane. PLoS One 16(5):e0245710. doi: 10.1371/journal.pone.0245710

Perreau J, Patel DJ, Anderson H, Maeda GP, Elston KM, Barrick JE, Moran NA. 2021. Vertical transmission at the pathogen-symbiont interface: Serratia symbiotica and aphids. mBio 12(2):e00359-21. doi: 10.1128/mBio.00359-21

Hammer TJ, Le E, Moran NA. 2021. Thermal niches of specialized gut symbionts: the case of social bees. Proc Biol Sci. 288(1944): 20201480. doi: 10.1098/rspb.2020.1480


Elston KM, Perreau J, Maeda GP, Moran NA, Barrick JE. 2020. Engineering a culturable Serratia symbiotica strain for aphid paratransgenesis. Appl Environ Microbiol. 87(4):e02245-20. doi: 10.1128/AEM.02245-20.

Horak RD, Leonard SP, Moran NA. 2020. Symbionts shape host innate immunity in honeybees. Proc Biol Sci. 287(1933):20201184. doi: 10.1098/rspb.2020.1184.

Motta EVS, Moran NA. 2020. Impact of Glyphosate on the Honey Bee Gut Microbiota: Effects of Intensity, Duration, and Timing of Exposure. mSystems 5(4):e00268-20. doi: 10.1128/mSystems.00268-20.

Motta EVS, Mak M, De Jong TK, Powell JE, O'Donnell A, Suhr KJ, Riddington IM, Moran NA. 2020. Oral and topical exposure to glyphosate in herbicide formulation impact the gut microbiota and survival rates of honey bees. Appl Environ Microbiol. 86(18):e01150-20. doi: 10.1128/AEM.01150-20.

Li Y, Zhang B, Moran NA. 2020. The aphid X chromosome is a dangerous place for functionally important genes: Diverse evolution of Hemipteran genomes based on chromosome-level assemblies. Mol Biol Evol. 37: 2357-2368. doi: 10.1093/molbev/msaa095.

Leonard SP, Powell JE, Perutka J, Geng P, Heckmann LC, Horak RD, Davies BW, Ellington AD, Barrick JE, Moran NA. 2020. Engineered symbionts activate honey bee immunity and limit pathogens. Science 367: 573-576. doi: 10.1126/science.aax9039 perspective
abstract | full text | reprint

Smith TE, Moran NA. 2020. Coordination of host and symbiont gene expression reveals a metabolic tug-of-war between aphids and Buchnera. Proc Natl Acad Sci U S A. 117: 2113-2121. doi: 10.1073/pnas.1916748117.


Zheng H, Perreau J, Powell JE, Han B, Zhang Z, Kwong WK, Tringe SG, Moran NA. 2019. Division of labor in honey bee gut microbiota for plant polysaccharide digestion. Proc Natl Acad Sci U S A. 116: 25909-25916. doi: 10.1073/pnas.1916224116

Zhang B, Leonard SP, Li Y, NA Moran. 2019. Obligate bacterial endosymbionts limit thermal tolerance of insect host species. Proc Natl Acad Sci U S A.116: 24712-24718. doi: 10.1073/pnas.1915307116.

Moran NA, Ochman H, Hammer TJ. 2019. Evolutionary and Ecological Consequences of Gut Microbial Communities. Annu Rev Ecol Syst. Vol. 50: 451-475. doi: 10.1146/annurev-ecolsys-110617-062453

Hammer TJ, Moran NA. 2019. Links between metamorphosis and symbiosis in holometabolous insects. Philos Trans R Soc Lond B Biol Sci. 374:20190068. doi: 10.1098/rstb.2019.0068.

Li Y, Park H, Smith TE, Moran NA. 2019. Gene family evolution in the pea aphid based on chromosome-level genome assembly. Mol Biol Evol. 36: 2143-2156. doi: 10.1093/molbev/msz138

Chong RA, Park H, Moran NA. 2019. Genome evolution of the obligate endosymbiont Buchnera aphidicola. Mol Biol Evol. 36:1481-1489. doi: 10.1093/molbev/msz082


Zheng H, Steele MI, Leonard SP, Motta EVS, Moran NA. 2018. Honey bees as models for gut microbiota research. Lab Anim (NY). 47: 317-325. doi: 10.1038/s41684-018-0173-x.

Raymann K, Coon KL, Shaffer Z, Salisbury S, Moran NA. 2018. Pathogenicity of Serratia marcescens Strains in Honey Bees. MBio. 9(5). pii: e01649-18. doi: 10.1128/mBio.01649-18.

Motta EVS, Raymann K, Moran NA. 2018. Glyphosate perturbs the gut microbiota of honey bees. Proc Natl Acad Sci U S A. 15: 10305-10310. doi: 10.1073/pnas.1803880115

Powell JE, Eiri D, Moran NA, Rangel J. 2018. Modulation of the honey bee queen microbiota: Effects of early social contact. PLoS One. 13(7) :e0200527. doi: 10.1371/journal.pone.0200527

Raymann K, Moran NA. 2018. The role of the gut microbiome in health and disease of adult honey bee workers. Curr Opin Insect Sci. 26: 97-104. doi: 10.1016/j.cois.2018.02.012

Raymann K, Motta EVS, Girard C, Riddington IM, Dinser JA, Moran NA. 2018. Imidacloprid decreases honey bee survival but does not affect the gut microbiome. Appl Environ Microbiol. 84(13). pii: e00545-18. doi: 10.1128/AEM.00545-18

Leonard SP, Perutka J, Powell JE, Geng P, Richhart DD, Byrom M, Kar S, Davies BW, Ellington AD, Moran NA, Barrick JE. 2018. Genetic engineering of bee gut microbiome bacteria with a toolkit for modular assembly of broad-host-range plasmids. ACS Synth Biol. 7: 1279-1290. doi: 10.1021/acssynbio.7b00399.

Kwong WK, Steele MI, Moran NA. 2018. Genome sequences of Apibacter spp., gut symbionts of Asian honey bees. Genome Biol Evol. 10: 1174-1179. doi: 10.1093/gbe/evy076.

Mockler BK, Kwong WK, Moran NA, Koch H. 2018. Microbiome structure influences infection by the parasite Crithidia bombi in bumble bees. Appl Environ Microbiol. 284(7). pii: e02335-17. doi: 10.1128/AEM.02335-17

Chong RA, Moran NA. 2018. Evolutionary loss and replacement of Buchnera, the obligate endosymbiont of aphids. ISME J. 12: 898-908. doi: 10.1038/s41396-017-0024-6

Raymann K, Bobay LM, Moran NA. 2018. Antibiotics reduce genetic diversity of core species in the honeybee gut microbiome. Mol Ecol. 27: 2057-2066. doi: 10.1111/mec.14434.


Steele MI, Kwong WK, Whiteley M, Moran NA. 2017. Diversification of type VI secretion system toxins reveals ancient antagonism among bee gut microbes. MBio. 8: e01630-17. doi: 10.1128/mBio.01630-17

Martinson VG, Carpinteyro-Ponce J, Moran NA, Markow TA. 2017. A distinctive and host-restricted gut microbiota in populations of a cactophilic Drosophila. Appl Environ Microbiol. 83: pii: e01551-17. doi: 10.1128/AEM.01551-17

Kwong WK, Zheng H, Moran NA. 2017. Convergent evolution of a modified, acetate-driven TCA cycle in bacteria. Nat Microbiol. 2:17067. doi: 10.1038/nmicrobiol.2017.67.

Zheng H, Powell JE, Steele MI, Dietrich C, Moran NA. 2017. Honeybee gut microbiota promotes host weight gain via bacterial metabolism and hormonal signaling. Proc Natl Acad Sci U S A. 114: 4775-4780. doi: 10.1073/pnas.1701819114

Kwong WK, Medina LA, Koch H, Sing K-W, Soh EJY, Ascher JS, Jaffé R, Moran NA. 2107. Dynamic microbiome evolution in social bees. Sci Adv. 3(3): e1600513. doi: 10.1126/sciadv.1600513

Raymann K, Shaffer Z, Moran NA. 2017. Antibiotic exposure perturbs the gut microbiota and elevates mortality in honeybees. PLoS Biol. 15(3):e2001861. doi: 10.1371/journal.pbio.2001861

Kwong WK, Mancenido AL, Moran NA. 2017. Immune system stimulation by the native gut microbiota of honey bees. R Soc Open Sci. 4: 170003.

Moran NA. 2017. Old and new symbiotic partners in lachnine aphids. Environ Microbiol. 19: 7. doi: 10.1111/1462-2920.13516. Commentary


Neuvonen MM, Tamarit D, Näslund K, Liebig J, Feldhaar H, Moran NA, Guy L, Andersson SG. 2016. The genome of Rhizobiales bacteria in predatory ants reveals urease gene functions but no genes for nitrogen fixation. Sci Rep. 6:39197. doi: 10.1038/srep39197.

Moran NA. 2016. When Obligate Partners Melt Down. MBio. 7(6). pii: e01904-16. doi: 10.1128/mBio.01904-16. Commentary

Powell JE, Leonard SP, Kwong WK, Engel P, Moran NA. 2016. Genome-wide screen identifies host colonization determinants in a bacterial gut symbiont. Proc Natl Acad Sci U S A.113:13887-13892. doi: 10.1073/pnas.1610856113

Chong RA, Moran NA. 2016. Intraspecific genetic variation in hosts affects regulation of obligate heritable symbionts. Proc Natl Acad Sci U S A. 113: 13114-13119/ doi: 10.1073/pnas.1610749113

Zheng H, Nishida A, Kwong WK, Koch H, Engel P, Steele MI, Moran NA. 2016. Metabolism of toxic sugars by strains of the bee gut symbiont Gilliamella apicola. mBio vol. 7 no. 6 e01326-16. doi: 10.1128/mBio.01326-16

Moran NA. 2016. Insights into the roles of bacterial symbionts within flagellates of termite guts. Environ Microbiol Rep. 2016 Sep 8. doi: 10.1111/1758-2229.12471. [Epub ahead of print] Commentary

Elijah Powell J, Ratnayeke N, Moran NA. 2016. Strain diversity and host specificity in a specialized gut symbiont of honey bees and bumble bees. Mol Ecol. 25: 4461-71. doi: 10.1111/mec.13787

Schwarz RS, Moran NA, Evans JD. 2016. Early gut colonizers shape parasite susceptibility and microbiota composition in honey bee workers. Proc Natl Acad Sci U S A. 113: 9345-9350. doi: 10.1073/pnas.1606631113

Tian B, Moran NA. 2016. Genome Sequence of Hafnia alvei bta3_1, a Bacterium with Antimicrobial Properties Isolated from Honey Bee Gut. Genome Announc. 4(3). pii: e00439-16. doi: 10.1128/genomeA.00439-16.

Engel P, Kwong WK, McFrederick Q, Anderson KE, Barribeau SM, Chandler JA, Cornman RS, Dainat J, de Miranda JR, Doublet V, Emery O, Evans JD, Farinelli L, Flenniken ML, Granberg F, Grasis JA, Gauthier L, Hayer J, Koch H, Kocher S, Martinson VG, Moran N, Munoz-Torres M, Newton I, Paxton RJ, Powell E, Sadd BM, Schmid-Hempel P, Schmid-Hempel R, Song SJ, Schwarz RS, vanEngelsdorp D, Dainat B. 2016. The bee microbiome: impact on bee health and model for evolution and ecology of host-microbe interactions. mBio 7(2):e02164-15. doi:10.1128/mBio.02164-15.

Kwong WK, Moran NA. 2016. Gut microbial communities of social bees. Nat Rev Microbiol. 14: 374-384. doi: 10.1038/nrmicro.2016.43

Kwong WK, Moran NA. 2016. Apibacter adventoris gen. nov., sp. nov., a member of the phylum Bacteroidetes isolated from honey bees. Int J Syst Evol Microbiol. 66: 1323-1329. doi: 10.1099/ijsem.0.000882


Moran NA, Sloan DB. 2015. The Hologenome Concept: Helpful or Hollow? PLoS Biol. 2015 Dec 4;13(12):e1002311. doi: 10.1371/journal.pbio.1002311

Bennett GM, McCutcheon JP, Mcdonald BR, Moran NA. 2015. Lineage-specific patterns of genome deterioration in obligate symbionts of sharpshooter leafhoppers. Genome Biol Evol. 8: 296-301. doi: 10.1093/gbe/evv159.

Li J, Powell JE, Guo J, Evans JD, Wu J, Williams P, Lin Q, Moran NA, Zhang Z. 2015. Two gut community enterotypes recur in diverse bumblebee species. Curr Biol. 25: R652-3. doi: 10.1016/j.cub.2015.06.031.

Moran NA. 2015. Genomics of the honey bee microbiome. Curr Opin Insect Sci. 10:22–28. doi: doi:10.1016/j.cois.2015.04.003

Kwong WK, Moran NA. 2015. Evolution of host specialization in gut microbes: the bee gut as a model. Gut Microbes. 6: 214-220. doi: 10.1080/19490976.2015.1047129

Engel P, Bartlett KD, Moran NA. 2015. The bacterium Frischella perrara causes scab formation in the gut of its honeybee host. MBio 6(3). pii: e00193-15. doi: 10.1128/mBio.00193-15.

Bennett GM, Moran NA. 2015. Heritable symbiosis: The advantages and perils of an evolutionary rabbit hole. Proc Natl Acad Sci U S A. 112: 10169-10176. doi: 10.1073/pnas.1421388112

Moran NA, Yun Y. 2015. Experimental replacement of an obligate insect symbiont. Proc Natl Acad Sci U S A. 112: 2093-2096. doi: 10.1073/pnas.1420037112


Kwong WK, Mancenido AL, Moran NA. 2014. Genome Sequences of Lactobacillus sp. Strains wkB8 and wkB10, Members of the Firm-5 Clade, from Honey Bee Guts. Genome Announc. 2(6) pii: e01176-14. doi: 10.1128/genomeA.01176-14.

Bennett GM, McCutcheon JP, MacDonald BR, Romanovicz D, Moran NA. 2014. Differential genome evolution between companion symbionts in an insect-bacterial symbiosis. MBio 5(5) pii: e01697-14. doi: 10.1128/mBio.01697-14.

Powell JE, Martinson VG, Urban-Mead K, Moran NA. 2014. Routes of acquisition of the gut microbiota of Apis mellifera. Appl Environ Microbiol. 80: 7378-7387. doi: 10.1128/AEM.01861-14.

Engel P, Stepanauskas R, Moran NA. 2014. Hidden diversity in honey bee gut symbionts detected by single-cell genomics. PLoS Genet. 10(9):e1004596. doi: 10.1371/journal.pgen.1004596.

Kwong WK, Engel P, Koch H, Moran NA. 2014. Genomics and host specialization of honey bee and bumble bee gut symbionts. Proc Natl Acad Sci U S A. 111: 11509-11514. doi: 10.1073/pnas.1405838111

Moran NA, Bennett GM. 2014. The tiniest tiny genomes. Annu Rev Microbiol. 68: 195-215. doi: 10.1146/annurev-micro-091213-112901.

Cariveau DP, Elijah Powell J, Koch H, Winfree R, Moran NA. 2014. Variation in gut microbial communities and its association with pathogen infection in wild bumble bees (Bombus). ISME J. 8(12):2369-2379. doi: 10.1038/ismej.2014.68

Martinson VG, Magoc T, Koch H, Salzberg SL, Moran NA. 2014. Genomic features of a bumble bee symbiont reflect its host environment. Appl Environ Microbiol. 80: 3793-3803. doi: 10.1128/AEM.00322-14

Sabree ZL, Moran NA. 2014. Host-specific assemblages typify gut microbial communities of related insect species. Springerplus 3:138. doi: 10.1186/2193-1801-3-138

Koga R, Moran NA. 2014. Swapping symbionts in spittlebugs: evolutionary replacement of a reduced genome symbiont. ISME J. 8: 1237-1246. doi: 10.1038/ismej.2013.235

Sloan DB, Nakabachi A, Richards S, Qu J, Murali SC, Gibbs RA, Moran NA. 2014. Parallel histories of horizontal gene transfer facilitated extreme reduction of endosymbiont genomes in sap-feeding insects. Mol Bio Evol. 31: 857-871. doi: 10.1093/molbev/msu004

Hansen AK, Moran NA. 2014. The impact of microbial symbionts on host plant utilization by herbivorous insects. Mol Ecol. 23: 1473-1496. doi: 10.1111/mec.12421.


Engel P, James R, Koga R, Kwong WK, McFrederick Q, Moran NA. 2013. Standard methods for research on Apis mellifera gut symbionts. In V Dietemann; J D Ellis; P Neumann (Eds) The COLOSS BEEBOOK, Volume I: standard methods for Apis mellifera research. Journal of Apicultural Research 52(4):

Bennett GM, Moran NA. 2013. Small, smaller, smallest: the origins and evolution of ancient dual symbioses in a phloem-feeding insect. Genome Biol Evol. 5: 1675-1688. doi: 10.1093/gbe/evt118

Engel P, Moran NA. 2013. The gut microbiota of insects - diversity in structure and function. FEMS Microbiol Rev. 37: 699-735. doi: 10.1111/1574-6976.12025.

Engel P, Kwong WK, Moran NA. 2013. Frischella perrara gen. nov., sp. nov., a gammaproteobacterium isolated from the gut of the honey bee, Apis mellifera. Int J Syst Evol Microbiol. 63: 3646-3651. doi: 10.1099/ijs.0.049569-0

Vogel KJ, Moran NA. 2013. Functional and evolutionary analysis of the genome of an obligate fungal symbiont. Genome Biol Evol. 5: 891-904. doi: 10.1093/gbe/evt054

Sloan DB, Moran NA. 2013. The evolution of genomic instability in the obligate endosymbionts of whiteflies. Genome Biol Evol. 5: 783-93. doi: 10.1093/gbe/evt044

Nováková E, Hypša V, Klein J, Foottit RG, Dohlen CD, Moran NA. 2013. Reconstructing the phylogeny of aphids (Hemiptera: Aphididae) using DNA of the obligate symbiont Buchnera aphidicola. Mol Phylogenet Evol. 68: 42-54. doi: 10.1016/j.ympev.2013.03.016

Koga R, Bennett G, Cryan JR, Moran NA. 2013. Evolutionary replacement of obligate symbionts in an ancient and diverse insect lineage. Environ Microbiol. 5: 2073-2081. doi: 10.1111/1462-2920.12121

<>Engel P, Moran NA. 2013. Functional and evolutionary insights into the simple yet specific gut microbiota of the honey bee from metagenomic analysis. Gut Microbes 4: 1-6. doi: 10.4161/gmic.22517

Kwong WK, Moran NA. 2013. Cultivation and characterization of the gut symbionts of honey bees and bumble bees: description of Snodgrassella alvi gen. nov., sp. nov., a member of the family Neisseriaceae of the Betaproteobacteria, and Gilliamella apicola gen. nov., sp. nov., a member of Orbaceae fam. nov., Orbales ord. nov., a sister taxon to the order 'Enterobacteriales' of the Gammaproteobacteria. Int J Syst Evol Microbiol. 63: 2008-2018. doi: 10.1099/ijs.0.044875-0

Sabree ZL, Huang CY, Okusu A, Moran NA, Normark BB. 2013. The nutrient supplying capabilities of Uzinura, an endosymbiont of armoured scale insects. Environ Microbiol. 15: 1988-1999. doi: 10.1111/1462-2920.12058.


Tian B, Fadhil NH, Powell JE, Kwong WK, Moran NA. 2012. Long-term exposure to antibiotics has caused accumulation of resistance determinants in the gut microbiota of honeybees. mBio 3(6):e00377-12. doi:10.1128/mBio.00377-12
Honeybees harbor antibiotic-resistance genes at ASM

Sloan DB, Moran NA. 2012. Endosymbiotic bacteria as a source of carotenoids in whiteflies. Biol Lett. 8: 986-989. doi: 10.1098/rsbl.2012.0664

Huang CY, Sabree ZL, Moran NA. 2012. Genome Sequence of Blattabacterium sp. Strain BGIGA, Endosymbiont of the Blaberus giganteus Cockroach. J Bacteriol. 194: 4450-4451. doi:10.1128/JB.00789-12

Sloan DB, Moran NA. 2012. Genome reduction and co-evolution between the primary and secondary bacterial symbionts of psyllids. Mol Bio Evol. 29: 3781-3792. doi: 10.1093/molbev/mss180

Sabree ZL, Hansen AK, Moran NA. 2012. Independent Studies Using Deep Sequencing Resolve the Same Set of Core Bacterial Species Dominating Gut Communities of Honey Bees. PLoS ONE 7(7): e41250. doi:10.1371/journal.pone.0041250

Engel P, Martinson VG, Moran NA. 2012. Functional diversity within the simple gut microbiota of the honey bee. Proceedings of the National Academy of Sciences USA. 109: 11002-11007. doi: 10.1073/pnas.1202970109
Editor's choice: Simple but specific in Science

Moran NA. 2012. Microbial symbiosis and evolution. pages 191-196 in Microbes and Evolution: The World that Darwin Never Saw, edited by R. Kolter and S. Maloy, ASM Press.

Hansen AK, Moran NA. 2012. Altered tRNA characteristics and 3' maturation in bacterial symbionts with reduced genomes. Nucleic Acids Res. 40: 7870–7884. doi: 10.1093/nar/gks503

Moran NA, Hansen AK, Powell JE, Sabree ZL. 2012. Distinctive Gut Microbiota of Honey Bees Assessed Using Deep Sampling from Individual Worker Bees. PLoS ONE 7(4): e36393. doi:10.1371/journal.pone.0036393

Martinson VG, Moy J, Moran NA. 2012.Establishment of characteristic gut bacteria during development of the honey bee worker. 2012. Appl. Environ. Microbiol. 78: 2830-2840. doi: 10.1128/AEM.07810-11

Hansen AK, Vorburger C, Moran NA. 2012. Genomic basis of endosymbiont-conferred protection against an insect parasitoid. Genome Res. 22:106-114. doi: 10.1101/gr.125351.111

Sabree ZL, Huang CY, Arakawa G, Tokuda G, Lo N, Watanabe H, Moran NA. 2012. Genome shrinkage and loss of nutrient-providing potential in the obligate symbiont of the primitive termite Mastotermes darwiniensis. Appl Environ Microbiol. 78: 204-210. doi: 10.1128/ AEM.06540-11

Nováková E, Moran NA. 2012. Diversification of genes for carotenoid biosynthesis in aphids following an ancient transfer from a fungus. Mol Bio Evol. 29: 313-323. doi: 10.1093/molbev/msr206


McCutcheon JP, Moran NA. 2011. Extreme genome reduction in symbiotic bacteria. Nat Rev Microbiol. 10: 13-26. doi: 10.1038/nrmicro2670.

Degnan PH, Ochman, H, Moran NA. 2011. Sequence conservation and functional constraint on intergenic spacers in reduced genomes of the obligate symbiont Buchnera. PLoS Genet. 7(9): e1002252. doi:10.1371/journal.pgen.1002252

Vogel KJ, Moran NA. 2011. Effect of host genotype on symbiont titer in the aphid—Buchnera symbiosis. Insects 2: 423-434; doi:10.3390/insects2030423

Burke GR, Moran NA. 2011. Responses of the pea aphid transcriptome to infection by facultative symbionts. Insect Mol Biol. 20: 357-365. doi: 10.1111/j.1365-2583.2011.01070.x

Degnan PH, Bittleston LS, Hansen AK, Sabree ZL, Moran NA, Almeida RP. 2011. Origin and examination of a leafhopper facultative endosymbiont. Curr Microbiol. 65: 1565-1572. doi: 10.1007/s00284-011-9893-5

Hansen AK, Moran NA. 2011. Aphid genome expression reveals host-symbiont cooperation in the production of amino acids. Proceedings of the National Academy of Sciences USA. 108: 2849-2854. doi: 10.1073/pnas.1013465108

Burke GR, Moran NA. 2011. Massive genomic decay in Serratia symbiotica, a recently evolved symbiont of aphids. Genome Biol Evol. 3: 195-208. doi: 10.1093/gbe/evr002

Martinson VG, Danforth BN, Minckley RL, Rueppell O, Tingek S, Moran NA. 2011. A simple and distinctive microbiota associated with honey bees and bumble bees. Molecular Ecology. 20: 619-628. doi: 10.1111/j.1365-294X.2010.04959.x
Commentary by Martin Kaltenpoth

Vogel KJ, Moran NA. 2011. Sources of variation in dietary requirements in an obligate nutritional symbiosis. Proceedings of the Royal Society: Biological Sciences. 278: 115-121. doi: 10.1098/rspb.2010.1304


McCutcheon JP, Moran NA. 2010. Functional convergence in reduced genomes of bacterial symbionts spanning 200 million years of evolution. Genome Biology and Evolution. 2: 708-718. doi: 10.1093/gbe/evq055 | Public access: Full Text or pdf

Burke GR, McLaughlin HJ, Simon JC, Moran NA. 2010. Dynamics of a recurrent Buchnera mutation that affects thermal tolerance of pea aphid hosts. Genetics 186(1): 367-72. doi:10.1534/genetics.110.117440

Sabree, ZL, Degnan PH, Moran NA. 2010. Chromosome stability and gene loss in cockroach endosymbionts. Applied and Environmental Microbiology 76(12):4076-9. doi:10.1128/AEM.00291-10

Woyke T, Tighe D, Mavromatis K, Clum A, Copeland A, Schackwitz W, Lapidus A, Wu D, McCutcheon JP, McDonald BR, Moran NA, Bristow J, Cheng JF. 2010. One bacterial cell, one complete genome. PLoS One 5: e10314. doi: 10.1371/journal.pone.0010314

Moran NA, Jarvik T. 2010. Lateral transfer of genes from fungi underlies carotenoid production in aphids. Science 328: 624-627. DOI: 10.1126/science.1187113 | Public access: Full Text or pdf
A Fungal Past to Insect Color in Science

Nikoh N, McCutcheon JP, Kudo T, Miyagishima SY, Moran NA, Nakabachi A. 2010. Bacterial genes in the aphid genome: absence of functional gene transfer from Buchnera to its host. PLoS Genetics 6: e1000827. doi: 10.1371/journal.pgen.1000827

International Aphid Genomics Consortium. 2010. Genome sequence of the pea aphid Acyrthosiphon pisum. PLoS Biology 8: e1000313. doi: 10.1371/journal.pbio.1000313

Oliver KM, Degnan PH, Burke GR, Moran NA. 2010. Facultative symbionts of aphids and the horizontal transfer of ecologically important traits. Annual Review of Entomology 55: 247-266. DOI: 10.1146/annurev-ento-112408-085305


Burke G, Fiehn O, Moran N. 2009. Effects of facultative symbionts and heat stress on the metabolome of pea aphids. The ISME Journal 4: 242-252. doi:10.1038/ismej.2009.114
Some species could be cushioned against climate change at UA News

Sabree ZL, Kambhampati S, Moran NA. 2009. Nitrogen recycling and nutritional provisioning by Blattabacterium, the cockroach endosymbiont. Proceedings of the National Academy of Sciences USA 106: 19521-19526. doi: 10.1073/pnas.0907504106

Degnan PH, Leonardo TE, Cass BN, Hurwitz B, Stern D, Gibbs RA, Richards S, Moran NA. 2009. Dynamics of genome evolution in facultative symbionts of aphids. Environmental Microbiology 12: 2060-2069. DOI: 10.1111/j.1462-2920.2009.02085.x

Peccoud J, Simon JC, McLaughlin HJ, Moran NA. 2009. Post-Pleistocene radiation of the pea aphid complex revealed by rapidly evolving endosymbionts. Proceedings of the National Academy of Sciences USA 106: 16315-16320. doi: 10.1073/pnas.0905129106

McCutcheon JP, McDonald BR, Moran NA. 2009. Convergent evolution of metabolic roles in bacterial co-symbionts of insects. Proceedings of the National Academy of Sciences USA 106: 15394-15399. doi: 10.1073/pnas.0906424106

Oliver KM, Degnan PH, Hunter MS, Moran NA. 2009. Bacteriophages encode factors required for protection in a symbiotic mutualism. Science 325: 992-994. DOI: 10.1126/science.1174463
Good Guys, Bad Guys on Small Things Considered

Nováková E, Hypsa V, Moran NA. 2009. Arsenophonus, an emerging clade of intracellular symbionts with a broad host distribution. BMC Microbiology 9: 143. doi:10.1186/1471-2180-9-143

McCutcheon JP, McDonald BR, Moran NA. 2009. Origin of an alternative genetic code in the extremely small and GC-rich genome of a bacterial symbiont. PLoS Genetics 5: e1000565. doi: 10.1371/journal.pgen.1000565
An Iconoclastic Endosymbiont on Small Things Considered

Burke GR, Normark BB, Favret C, Moran NA. 2009. Evolution and diversity of facultative symbionts from the aphid subfamily Lachninae. Applied Environmental Microbiology 75: 5328-35. doi:10.1128/AEM.00717-09

Kuo CH, Moran NA, Ochman H. 2009. The consequences of genetic drift for bacterial genome complexity. Genome Research 19: 1450-4. doi: 10.1101/gr.091785.109

Watts T, Haselkorn TS, Moran NA, Markow TA. 2009. Variable incidence of Spiroplasma infections in natural populations of Drosophila species. PLoS One 4: e5703. doi: 10.1371/journal.pone.0005703

Degnan PH, Yu Y, Sisneros N, Wing RA, Moran NA. 2009. Hamiltonella defensa, genome evolution of protective bacterial endosymbiont from pathogenic ancestors. Proceedings of the National Academy of Sciences USA 106: 9063-9068. doi: 10.1073/pnas.0900194106

Harmon JP, Moran NA, Ives AR. 2009. Species response to environmental change: impacts of food web interactions and evolution. Science 323(5919): 1347-50. doi: 10.1126/science.1167396

Haselkorn TS, Markow TA, Moran NA. 2009. Multiple introductions of the Spiroplasma bacterial endosymbiont into Drosophila. Molecular Ecology 18(6): 1294-305. doi: 10.1111/j.1365-294X.2009.04085.x

Moran NA, McLaughlin HJ, Sorek R. 2009. The dynamics and time scale of ongoing genomic erosion in symbiotic bacteria. Science. 323: 379-382. doi: 10.1126/science.1167140
Sequencing bacteria gives us a look at genomic erosion (Science 2.0)


Moran NA, McCutcheon JP, Nakabachi A. 2008. Genomics and evolution of heritable bacterial symbionts. Annual Review of Genetics 42: 165-190. doi: 10.1146/annurev.genet.41.110306.130119

Degnan PH, Moran NA. 2008. Diverse phage-encoded toxins in a protective insect endosymbiont. Applied Environmental Microbiology 74(21): 6782-91. doi:10.1128/AEM.01285-08

Degnan PH, Moran NA. 2008. Evolutionary genetics of a defensive facultative symbiont of insects: exchange of toxin-encoding bacteriophage. Molecular Ecology 17(3): 916-29. doi: 10.1111/j.1365-294X.2007.03616.x

Oliver KM, Campos J, Moran NA, Hunter MS. 2008. Population dynamics of defensive symbionts in aphids. Proceedings of the Royal Society B 275(1632): 293-9. doi: 10.1098/rspb.2007.1192

Plague GR, Dunbar HE, Tran PL, Moran NA. 2008. Extensive proliferation of transposable elements in heritable bacterial symbionts. Journal of Bacteriology 190(2): 777-9. doi:10.1128/JB.01082-07


McCutcheon JP, Moran NA. 2007. Parallel genomic evolution and metabolic interdependence in an ancient symbiosis. Proceedings of the National Academy of Sciences USA 104: 19392-7. doi: 10.1073/pnas.0708855104
It Takes Three to Tango on Small Things Considered

Cox-Foster DL, Conlan S, Holmes EC, Palacios G, Evans JD, Moran NA,  Quan PL, Briese T, Hornig M, Geiser DM, Martinson V, vanEngelsdorp D, Kalkstein AL, Drysdale A, Hui J, Zhai J, Cui L, Hutchison SK, Simons JF, Egholm M, Pettis JS, Lipkin WI. 2007. A metagenomic survey of microbes in honey bee colony collapse disorder. Science 318(5848): 283-7. doi: 10.1126/science.1146498

Moran NA. 2007. Symbiosis as an adaptive process and source of phenotypic complexity. Proceedings of the National Academy of Sciences USA 104 Suppl 1: 8627-33. doi: 10.1073/pnas.0611659104

Dunbar HE, Wilson AC, Ferguson NR, Moran NA. 2007. Aphid thermal tolerance is governed by a point mutation in bacterial symbionts. PLoS Biology 5(5): e96. doi: 10.1371/journal.pbio.0050096
When bacteria loose a single DNA base, aphids suffer at PLoS Biology


Moran NA. 2006. Symbiosis. Current Biology 16(20): R866-71. doi: 10.1016/j.cub.2006.09.019

Takiya DM, Tran P, Dietrich CH, Moran NA. 2006. Co-cladogenesis spanning three phyla: leafhoppers (Insecta: Hemiptera: Cicadellidae) and their dual bacterial symbionts. Molecular Ecology 15(13): 4175-91. doi: 10.1111/j.1365-294X.2006.03071.x

Nakabachi A, Yamashita A, Toh H, Ishikawa H, Dunbar HE, Moran NA, Hattori M. 2006. The 160-kilobase genome of the bacterial endosymbiont Carsonella. Science 314(5797): 267. doi: 10.1126/science.1134196
The Bacterium That Doesn't Know How To Tie Its Own Shoelaces on Small Things Considered

Moran NA, Dunbar HE. 2006. Sexual acquisition of beneficial symbionts in aphids. Proceedings of the National Academy of Sciences U S A 103(34): 12803-6. doi: 10.1073/pnas.0605772103

Dale C. Moran NA. 2006. Molecular interactions between bacterial symbionts and their hosts. Cell. 126(3): 453-65. doi: 10.1016/j.cell.2006.07.014

Mateos M, Castrezana SJ, Nankivell BJ, Estes A, Markow TA, Moran NA. 2006. Heritable endosymbionts of Drosophila. Genetics 174: 363–76. doi:10.1534/genetics.106.058818

Wu D, Daugherty SC, Van Aken SE, Pai GH, Watkins KL, Khouri H, Tallon LJ, Zaborsky JM, Dunbar HE, Tran PL, Moran NA, Eisen JA. 2006. Metabolic complementarity and genomics of the dual symbiosis of sharpshooters. PloS Biology 4(6): e188. doi: 10.137/journal.pbio.0040188

Oliver KM, Moran NA, Hunter MS. 2006. Costs and benefits of a superinfection of facultative symbionts in aphids. Proceedings of the Royal Society B 273(1591): 1273-80. doi: 10.1098/rspb.2005.3436

Moran NA, Degnan PH. 2006. Functional genomics of Buchnera and the ecology of aphid hosts. Molecular Ecology 15(5): 1251-61. doi: 10.1111/j.1365-294X.2005.02744.x

Russell JA, Moran NA. 2006. Costs and benefits of symbiont infection in aphids: variation among symbionts and across temperatures. Proceedings of the Royal Society B 273(1586): 603-10. doi: 10.1098/rspb.2005.3348

Wilson ACC, Dunbar HE, Davis GK, Hunter WB, Stern DL, Moran NA. 2006. A dual-genome microarray for the pea aphid, Acyrthosiphon pisum and its obligate bacterial symbiont, Buchnera aphidicola. BMC Genomics 7: 50. doi:10.1186/1471-2164-7-50


Moran NA, Tran P, Gerardo NM. 2005. Symbiosis and insect diversification: an ancient symbiont of sap-feeding insects from the bacterial phylum Bacteroidetes. Applied and Environmental Microbiology 71(12): 8802-10. doi:10.1128/AEM.71.12.8802-8810.2005

Russell JA, Moran NA. 2005. Horizontal transfer of bacterial symbionts: heritability and fitness effects in a novel aphid host. Applied and Environmental Microbiology 71(12): 7987-94. doi:10.1128/AEM.71.12.7987-7994.2005

Moran NA, Degnan PH, Santos SR, Dunbar HE, Ochman H. 2005. The players in a mutualistic symbiosis: Insects, bacteria, viruses and virulence genes. Proceedings of the National Academy of Sciences USA 102(47): 16919-26. doi: 10.1073/pnas.0507029102

Oliver KM, Moran NA, Hunter MS. 2005. Variation in resistance to parasitism in aphids is due to symbionts and not host genotype. Proceedings of the National Academy of Sciences USA 102(36): 12795-800. doi: 10.1073/pnas.0506131102
The right hitchhiker can save an aphid’s life at UA News

Moran NA, Dunbar HE, Wilcox JL. 2005. Regulation of transcription in a reduced bacterial genome: nutrient-provisioning genes of the obligate symbiont, Buchnera aphidicola. Journal of Bacteriology 187(12): 4229-37. doi:10.1128/JB.187.12.4229-4237.2005

Moran NA, Russell JA, Koga R, Fukatsu T. 2005. Evolutionary relationships of three new species of Enterobacteriaceae living as symbionts of aphids and other insects. Applied and Environmental Microbiology 71(6): 3302-10. doi:10.1128/AEM.71.6.3302-3310.2005

Lerat E, Daubin V, Ochman H, Moran NA. 2005. Evolutionary origins of genomic repertoires in bacteria. PLoS Biology 3(5): e130. doi: 10.1371/journal.pbio.0030130
Where bacteria get their genes at Science Daily


Dale CD, Dunbar HE, Moran NA, Ochman H. 2004. Extracting single genomes from heterogenous DNA samples: A test case with Carsonella ruddii the bacterial symbiont of psyllids (Insecta). Journal of Insect Science 5: 3.

Daubin V, Moran NA. 2004. Comment on The Evolution of Genome Complexity. Science 306: 978. doi: 10.1126/science.1098469

Lerat E, Moran NA. 2004. The evolutionary history of quorum-sensing systems in bacteria. Molecular Biology and Evolution 21: 903-913. doi: 10.1093/molbev/msh097

Moran NA, Plague GR. 2004. Genomic changes following host restriction in bacteria. Current Opinions in Genetics and Development 14: 627-633. doi: 10.1016/j.gde.2004.09.003


Dale C, Wang B, Moran N, Ochman H. 2003. Loss of DNA recombinational repair enzymes in the initial stages of genome degeneration in mutualistic bacterial endosymbionts. Molecular Biology and Evolution 20: 1188-1194. doi: 10.1093/molbev/msg138

Daubin V, Moran NA, Ochman H. 2003. Phylogenetics and the cohesion of bacterial genomes. Science 5634: 829-832. doi: 10.1126/science.1086568

Lerat E, Daubin V, Moran NA. 2003. From gene trees to organismal phylogeny in prokaryotes: the case of the γ-Proteobacteria. PLoS Biology 1: 101-108. doi: 10.1371/journal.pbio.0000019

Moran NA. 2003. Tracing the evolution of gene loss in obligate symbionts. Current Opinions in Microbiology 6: 512-518. doi:10.1016/j.mib.2003.08.001

Moran NA, Dale C, Dunbar H, Smith W, Ochman H. 2003. Intracellular symbionts of sharpshooters (Insecta: Hemiptera: Cicadellinae) form a distinct clade with a small genome. Environmental Microbiology 5: 116-126. doi: 10.1046/j.1462-2920.2003.00391.x

Moran NA, Plague G, Sandström J, Wilcox JL. 2003. A genomic perspective on nutrient-provisioning by bacterial symbionts of insects. Proceedings of the National Academy of Sciences USA (Colloquium Issue) 100: 14543-14548. doi: 10.1073/pnas.2135345100

Moran NA, Wilcox JL. 2003. Genomic evolution in Buchnera the obligate symbiont of aphids. Nova Acta Leopoldina 333:335-338.

Normark BB, Judson O, Moran NA. 2003. Genomic signatures of ancient asexual lineages. Biological Journal of the Linnean Society 79: 69-84. doi: 10.1046/j.1095-8312.2003.00182.x

Oliver KM, Russell JA, Moran NA, Hunter MS. 2003. Facultative bacterial symbionts in aphids confer resistance to parasitic wasps. Proceedings of the National Academy of Sciences USA 100:1803-1807. doi: 10.1073/pnas.0335320100

Plague GR, Dale C, Moran NA. 2003. Low and homogeneous copy number of plasmid-borne symbiont genes affecting host nutrition in Buchnera aphidicola of the aphid Uroleucon ambrosiae. Molecular Ecology 12: 1095-1100. doi: 10.1046/j.1365-294X.2003.01782.x

Russell JA, LaTorre AL, Sabater-Munoz B, Moya A, Moran NA. 2003. Side-stepping secondary symbionts: widespread horizontal transfer across and beyond the Aphidoidea. Molecular Ecology 12: 1061-1075. doi: 10.1046/j.1365-294X.2003.01780.x

Wilcox JL, Dunbar HE, Wolfinger RD, Moran NA. 2003. Consequences of reductive evolution for gene expression in an obligate endosymbiont. Molecular Microbiology 48: 1491-1500. doi: 10.1046/j.1365-2958.2003.03522.x


Abbot P, Moran NA. 2002. Extremely low levels of genetic polymorphism in endosymbionts of aphids (Pemphigus). Molecular Ecology 11: 2649-2660. doi: 10.1046/j.1365-294X.2002.01646.x

Dale C, Plague G, Wang B, Ochman H, Moran NA. 2002. Type III secretion and the conversion from parasitism to mutualism. Proceedings of the National Academy of Sciences USA 99: 12397-12402. doi: 10.1073/pnas.182213299

Mira A, Moran NA. 2002. Estimating population size and transmission bottlenecks in maternally transmitted endosymbiotic bacteria. Microbial Ecology 44: 137-143. doi: 10.1007/s00248-002-0012-9

Moran NA. 2002. The ubiquitous and varied role of infection in the lives of animals and plants. The American Naturalist 160:S1-S8. doi: 10.1086/342113

Moran NA. 2002. Microbial minimalism: Genome reduction in bacterial pathogens. Cell 108: 583-586. doi: 10.1016/S0092-8674(02)00665-7

Tamas I, Klasson L, Canbäck B, Näslund AK, Eriksson AS, Wernegreen JJ, Sandström JP, Moran NA, Andersson SG. 2002. Fifty million years of genomic stasis in endosymbiotic bacteria. Science 296:2376-2379. doi: 10.1126/science.1071278


Abbot P, Withgott JH, Moran NA. 2001. Genetic conflict and conditional altruism in social aphid colonies. Proceedings of the National Academy of Sciences USA 98:12068-12071. doi: 10.1073/pnas.201212698

Clark MA, Baumann L, Thao ML, Moran NA, Baumann P. 2001. Degenerative minimalism in the genome of a psyllid endosymbiont. Journal of Bacteriology 183: 1853-1861. doi: 10.1128/JB.183.6.1853-1861.2001

Funk DJ, Wernegreen JJ, Moran NA. 2001. Intraspecific variation in symbiont genomes: bottlenecks and the aphid- Buchnera association. Genetics 157: 477-489

Mira A, Ochman H, Moran NA. 2001. Deletional bias and the evolution of bacterial genomes. Trends in Genetics 17: 589-596. doi: 10.1016.S0168-9525(01)02447-7

Moran NA. 2001. Bacterial menageries inside insects. (Commentary). Proceedings of the National Academy of Sciences USA 98: 1338-1340. doi: 10.1073/pnas.98.4.1338

Moran NA. 2001. The coevolution of bacterial endosymbionts and phloem-feeding insects. Annals of the Missouri Botanical Gardens 88: 35-44.

Moran NA, Mira A. 2001. The process of genome shrinkage in the obligate symbiont, Buchnera aphidicola. Genome Biology 2: research0054.1-0054.12. doi:10.1186/gb-2001-2-12-research0054

Ochman H, Moran NA. 2001. Genes lost and genes found: the evolution of bacterial pathogenesis and symbiosis. Science 292: 1096-1098. doi: 10.1126/science.1058543

Sandström JP, Moran NA. 2001. Amino acid budgets in three aphid species using the same host plant. Physiological Entomology 26: 202-211. doi: 10.1046/j.0307-6962.2001.00235.x

Sandström JP, Russell JA, White JP, Moran NA. 2001. Independent origins and horizontal transfer of bacterial symbionts of aphids. Molecular Ecology 10: 217-228. doi: 10.1046/j.1365-294X.2001.01189.x

Thao ML, Clark MA, Burckhart DH, Moran NA, Baumann P. 2001. Phylogenetic analysis of vertically transmitted psyllid endsymbionts (Candidatus Carsonella ruddi) based on atpAGD and rpoC : Comparisons with 16S-23S rDNA-derived phylogeny. Current Microbiology 42: 419-421. doi: 10.1007/s002840010240

Wernegreen JJ, Moran NA. 2001. Vertical transmission of biosynthetic plasmids in aphid endosymbionts (Buchnera). Journal of Bacteriology 183: 785-790. doi: 10.1128/JB.183.2.785-790.2001

Wernegreen JJ, Richardson A, Moran NA. 2001. Parallel acceleration of evolutionary rates in genes underlying host nutrition in aphid symbionts. Molecular Phylogenetics and Evolution 19: 479-485. doi: 10.1006/mpev.2001.0929


Baumann P, Baumann L, Moran NA. 2000. Genus Buchnera Munson, Baumann, and Kinsey 1991. (entry in Bergey's Manual of Systematic Bacteriology).

von Dohlen CD, Moran NA. 2000. Molecular data support a rapid radiation of aphids in the Cretaceous and multiple origins of a complex life cycle. Biological Journal of the Linnean Society 71: 689-717. doi: 10.1111/j.1095-8312.2000.tb01286.x

Bernays EA, Funk DJ, Moran NA. 2000. Intraspecific differences in olfactory sensilla in relation to diet breadth in Uroleucon ambrosiae (Homoptera: Aphididae). Journal of Morphology 245: 99-109. doi: 10.1002/1097-4687(200008)245:2<99::AID-JMOR2>3.0.CO;2-O

Clark MA, Moran NA, Baumann P, Wernegreen JJ. 2000. Cospeciation between bacterial endosymbionts (Buchnera) and a recent radiation of aphids (Uroleucon) and pitfalls of testing for phylogenetic congruence. Evolution 54: 517-525. doi: 10.1111/j.0014-3820.2000.tb00054.x

Funk DJ, Helbling L, Wernegreen JJ, Moran NA. 2000. Intraspecific phylogenetic congruence among multiple symbiont genomes. Proceedings of the Royal Society of London Series B 267: 2517-2521. doi: 10.1098/rspb.2000.1314

Moran NA, Baumann P. 2000. Bacterial endosymbionts in animals. Current Opinions in Microbiology 3: 270-275. doi: 10.1016/S1369-5274(00)00088-6

Moran NA, Wernegreen JJ. 2000. Lifestyle evolution in symbiotic bacteria: insights from genomics. Trends in Ecology and Evolution 15: 321-326. doi 10.1016/S0169-5347(00)01902-9

Normark BB, Moran NA. 2000. Testing for the accumulation of deleterious mutations in asexual eukaryotes. Journal of Natural History 34:1719-1729. doi: 10.1080/00222930050122147

Rispe C, Moran NA. 2000. Accumulation of deleterious mutations in endosymbionts: Muller's ratchet with two levels of selection. The American Naturalist 156: 425-441. doi: 10.1086/303396

Sandström J, Telang A, Moran NA. 2000. Nutritional enhancement of host plants by aphids--a comparison of three aphid species on grasses. Journal of Insect Physiology 46: 33-40. doi: 10.1016/S0022-1910(99)00098-0

Thao ML, Clark M, Baumann L, Brennan EB, Moran NA, Baumann P. 2000. Secondary symbionts of psyllids have been acquired multiple times. Current Microbiology 41: 300-304. doi: 10.1007/s002840010138

Thao ML, Moran NA, Abbot P, Brennan EB, Burckhardt DH, Baumann P. 2000. Cospeciation of psyllids and their prokaryotic endosymbionts. Applied and Environmental Microbiology 66: 2898-2905.

Wernegreen JJ, Moran NA. 2000. Decay of mutualistic potential in aphid endosymbionts through silencing of biosynthetic loci: Buchnera of Diuraphis. Proceedings of the Royal Society Series B 267: 1423-1431. doi:10.1098/rspb.2000.1159

Wernegreen JJ, Ochman H, Jones I, Moran NA. 2000. The decoupling of genome size and sequence divergence in a symbiotic bacterium. Journal of Bacteriology 182: 3867-3869.


Baumann L, Baumann P, Moran NA, Sandström J, Thao ML. 1999. Genetic characterization of plasmids contining genes encoding enzymes of leucine biosynthesis in endosymbionts (Buchnera) of aphids. Journal of Molecular Evolution 48:77-85. doi: 10.1007/PL00006447

Baumann P, Moran NA,  Baumann L. 1999. Bacteriocyte-associated endosymbionts of insects. In The Prokaryotes, Third Edition, a Handbook on the Biology of Bacteria: Ecophysiology, isolation, identification, applications, (Dworkin, M., ed.) Springer-Verlag. PDF on ResearchGate

Clark MA, Baumann P, Moran NA. 1999. Buchnera plasmid-associated trpEG probably originated from a chromosomal location between hsIU and fpr. Current Microbiology 38: 309-311. doi: 10.1007/PL00006807

Clark MA, Moran NA, Baumann P. 1999. Sequence evolution in bacterial endosymbionts having extreme base composition. Molecular Biology and Evolution 16: 1586-1598.

Holt RD, McPeek MA, Moran NA. 1999. 1998 Sewall Wright Award: William Donald Hamilton. American Naturalist 153(1): i-ii. doi: 10.1086/303156

Moran NA, Kaplan MA, Gelsey Murphy MT, Scholes E. 1999. Phylogenetics and evolution of the aphid genus Uroleucon based on mitochondrial and nuclear DNA sequences. Systematic Entomology 24: 85-93. doi: 10.1046/j.1365-3113.1999.00076.x

Ochman H, Elwyn S, Moran NA. 1999. Calibrating bacterial evolution. Proceedings of the National Academy of Sciences USA 96:12638-12643 doi: 10.1073/pnas.96.22.12638

Sandström J, Moran NA. 1999. How nutritionally imbalanced is phloem sap for aphids? Entomologia experimentalis et applicata 91: 203-210. DOI: 10.1046/j.1570-7458.1999.00485.x

Telang A, Sandström J, Dyreson E, Moran NA. 1999. Feeding damage by Diuraphis noxia results in a nutritionally enhanced phloem diet. Entomologia experimentalis et applicata 91: 403-412. doi: 10.1046/j.1570-7458.1999.00508.x

Wernegreen J.J. and N. A. Moran. 1999. Evidence for genetic drift in endosymbionts: analyses of protein coding genes. Molecular Biology and Evolution 16: 83-97


Baumann L, Baumann P, Moran NA. 1998. The endosymbiont (Buchnera) of the aphid Diuraphis noxia contains all the genes of the tryptophan biosynthetic pathway. Current Microbiology 37:58-59. doi: 10.1007/s002849900337

Brynnel EA, Kurland CG, Andersson SGE, Moran NA. 1998. Evolutionary rates for tuf genes in endosymbionts of aphids. Molecular Biology and Evolution 15: 574-582.

Lambert JL, Moran NA. 1998. Deleterious mutations destabilize ribosomal RNA of endosymbionts. Proceedings of the National Academy of Sciences USA 95: 4458-4462.

Moran NA, Telang A. 1998. Bacteriocyte-associated endosymbionts in insects. Bioscience 48: 295-304.

Thao ML, Baumann L, Baumann P, Moran NA. 1998. Endosymbionts (Buchnera) from the aphids Schizaphis graminum and Diuraphis noxia have different copy numbers of the plasmid containing the leucine biosynthetic genes. Current Microbiology 36: 238-240. doi: 10.1007/s002849900301


Baumann L, Clark MA, Rouhbakhsh D, Baumann P, Moran NA, Voegtlin DJ. 1997. Endosymbionts (Buchnera) of Uroleucon sonchi contain plasmids with trpEG and remnants of trpE pseudogenes. Current Microbiology 35: 18- 21. doi: 10.1007/s002849900204

Baumann P, Moran NA. 1997. Non-cultivable microorganisms from symbiotic associations of insects and other hosts. Antonie van Leeuwenhoek 72: 39-48. doi: 10.1023/A:1000239108771

Baumann P, Moran NA, Baumann L. 1997. The evolution and genetics of aphid endosymbionts. Bioscience 47: 12-20.

Rouhbaksh D, Clark MA, Baumann L, Moran NA, Baumann P. 1997. Evolution of the tryptophan biosynthetic pathway in Buchnera (aphid endosymbionts): studies of plasmid-associated trpEG within the genus Uroleucon. Molecular Phylogenetics and Evolution 8: 167-176. doi: 10.1006/mpev.1997.0419

Withgott JH, Abbot DK, Moran NA. 1997. Maternal death relaxes developmental inhibition in nymphal aphid defenders. Proceedings of the Royal Society of London Series B 264: 1197-1202. doi: 10.1098/rspb.1997.0165


Aoki S, Kurosu U, Moran NA. 1996. Secondary monoecy of the gall aphid Thecabius populimonilis. Japanese Journal of Entomology 64: 367-378.

Lai C-Y, Baumann P, Moran NA. 1996. The endosymbiont (Buchnera) of the aphid Diuraphis noxia contains plasmids consisting of trpEG and tandem repeats of trpEG pseudogenes. Applied and Environmental Microbiology 62: 332-339.

Moran NA. 1996. Accelerated evolution and Muller's ratchet in endosymbiotic bacteria. Proceedings of the National Academy of Sciences 93: 2873-2878.

Rouhbaksh D, Lai C-Y, C. von Dohlen D, Baumann L, Baumann P, Moran NA, Voegtlin DJ. 1996. The tryptophan biosynthetic pathway of aphid endosymbionts (Buchnera): genetics and evolution of plasmid-associated trpEG within the Aphididae. Journal of Molecular Evolution 42: 414-421. doi: 10.1007/BF02498635

Seger J, Moran NA. 1996. Behavioural ecology - Snapping social swimmers. Nature 381: 473-474. doi:10.1038/381473a0


Baumann P, Baumann L, Lai C-Y, Rouhbakhsh D, Moran NA, Clark MA. 1995. Genetics, physiology, and evolutionary relationships of the genus Buchnera: intracellular symbionts of aphids. Annual Review of Microbiology 41: 55-94. doi: 10.1146/annurev.mi.49.100195.000415

Baumann P, Lai C-Y, Baumann L, Rouhbakhsh D, Moran NA, Clark MA. 1995. Mutualistic associations of aphids and prokaryotes: biology of the genus Buchnera. Applied and Environmental Microbiology 61: 1-7.

Lai C-Y, Baumann P, Moran NA. 1995. Genetics of the tryptophan biosynthetic pathway of the prokaryotic endosymbiont (Buchnera) of the aphid Schlechtendalia chinensis. Insect Molecular Biology 4: 47-59. doi: 10.1111/j.1365-2583.1995.tb00007.x

Moran NA, von Dohlen CD, Baumann P. 1995. Faster evolutionary rates in endosymbiotic bacteria than in cospeciating insect hosts. Journal of Molecular Evolution 41: 727-731. doi: 10.1007/BF00173152

von Dohlen CD, Moran NA. 1995. Molecular phylogeny of the Homoptera: a paraphyletic taxon. Journal of Molecular Evolution 41: 211-223. doi: 10.1007/BF00170675


Aoki S, Moran NA. 1994. Pemphigus obesinymphae, a new American aphid species with defenders and swollen nymphs (Homoptera: Aphidoidea: Pemphigidae). Journal of the New York Entomological Society 102: 251-260.

Moran NA. 1994. Adaptation and constraint in the complex life cycles of animals. Annual Review of Ecology and Systematics 25: 573-600. doi: 10.1146/

Moran NA, Baumann P. 1994. Phylogenetics of cytoplasmically inherited microorganisms of arthropods. Trends in Ecology and Evolution 9: 15-20. doi:10.1016/0169-5347(94)90226-7

Moran NA, Baumann P, von Dohlen C. 1994. Use of DNA sequences to reconstruct the history of the association between members of the Sternorrhyncha and their bacterial endosymbionts. European Journal of Entomology 91: 79-83.

Rouhbaksh D, Moran NA, Baumann L, Voegtlin DJ, Baumann P. 1994. Detection of Buchnera the primary prokaryotic endosymbiont of aphids, using the polymerase chain reaction. Insect Molecular Biology 3: 213-217. doi: 10.1111/j.1365-2583.1994.tb00169.x


Baumann P, Munson MA, Lai C-Y, Clark MA, Baumann L, Moran NA, Campbell BC. 1993. Origin and properties of bacterial endosymbionts of aphids, whiteflies, and mealybugs. American Society of Microbiology News 59: 21-24. PDF on ResearchGate

Moran NA. 1993. Defenders in the North American aphid, Pemphigus obesinymphae. Insectes Sociaux 40: 391-402. doi: 10.1007/BF01253902

Moran NA, Munson MA, Baumann P, Ishikawa H. 1993. A molecular clock in endosymbiotic bacteria is calibrated using the insect hosts. Proceedings of the Royal Society of London Series B 253: 167-171. doi: 10.1098/rspb.1993.0098

Moran NA, Seminoff J, Johnstone L. 1993. Genotypic variation in propensity for host alternation within a population of the aphid Pemphigus betae. Journal of Evolutionary Biology 6: 691-705. doi: 10.1046/j.1420-9101.1993.6050691.x

Moran NA, Seminoff J, Johnstone L. 1993. Induction of winged sexuparae in root- inhabiting colonies of the aphid Pemphigus betae. Physiological Entomology 18: 296-302. doi: 10.1111/j.1365-3032.1993.tb00602.


Clark MA, Baumann L, Munson MA, Baumann P, Campbell BC, Duffus JE, Osborne LS, Moran NA. 1992. The eubacterial endosymbionts of whiteflies (Homoptera: Aleyrodoidea) constitute a lineage distinct from the endosymbionts of aphids and mealybugs. Current Microbiology 25: 119-123. doi: 10.1007/BF01570970

Moran NA. 1992. Evolution of sex ratio variation in aphids. chapter in Evolution and Diversity of Sex Ratios in Insects and Mites (D. Wrensch and M. Ebbert, eds.), pp. 346-368. Chapman and Hall, NY.

Moran NA. 1992. The evolutionary maintenance of alternative phenotypes. The American Naturalist 139: 971-989.

Moran NA. 1992. The evolution of life cycles in aphids. Annual Review of Entomology 37: 321-348. doi: 10.1146/annurev.en.37.010192.001541

Munson M, Baumann P, Moran NA. 1992. Phylogenetic relationships of mealybug endosymbionts (Homoptera: Pseudococcidae) based on 16S rDNA sequences. Molecular Phylogenetics and Evolution 1: 26-30. doi:10.1016/1055-7903(92)90032-C


Moran NA. 1991. Phenotype fixation and genotypic diversity in the life cycle of the aphid, Pemphigus betae. Evolution 45: 957-970.

Munson MA, Baumann P, Clark MA, Baumann L, Moran NA, Voegtlin DJ, Campbell BC. 1991. Evidence for the establishment of aphid-eubacterium endosymbiosis in an ancestor of four aphid families. Journal of Bacteriology 173: 6321-6324.


Moran, N. A. 1990. Genetic variation in life cycle and host plant relations in Pemphigus betae. 7th International Symposium on Insect/Host Plant Relations. Symp. Biol. Hung. 39: 245-248.

Moran NA. 1990. Aphid life cycles: two evolutionary steps. The American Naturalist 136: 135-138.

Moran NA, Whitham TJ. 1990. Differential colonization of resistant and susceptible hostplants: Pemphigus and Populus. Ecology 71:1059-1067.

Moran NA, Whitham TG. 1990. Interspecific competition between root-feeding and leaf-galling aphids mediated by host plant resistance. Ecology 71: 1050-105.


Moran NA. 1989. A 48-million-year-old aphid-host plant association and complex life cycle: biogeographic evidence. Science 245: 173-175. doi: 10.1126/science.245.4914.173


Moran NA. 1988. The evolution of host alternation in aphids: evidence that specialization is a dead end. The American Naturalist 132: 681-706.

Moran NA, Whitham TG. 1988. Evolutionary reduction of complex life cycles: loss of host alternation in Pemphigus (Homoptera: Aphididae). Evolution 42: 717-728.

Moran NA, Whitham TG. 1988. Population fluctuations in complex life cycles: an example from Pemphigus aphids. Ecology 69: 1214-1218.


Moran NA. 1987. Evolutionary determinants of host plant specificity in Uroleucon. Pages 29-38 in Population Structure, Genetics, and Taxonomy of Aphids and Thysanoptera, edited by J. Holman, J. Pelikan, A. F. G. Dixon, and L. Weisman. SPB Publishing, The Hague, The Netherlands.


Moran NA. 1986. Morphological adaptation to host plants in Uroleucon (Homoptera: Aphididae). Evolution 40: 1044-1058.

Moran NA. 1986. Benefits of host plant specificity in Uroleucon (Homoptera: Aphididae). Ecology 67, 108-115.


Stenseth NC, Kirkendall LR, Moran NA. 1985. On the evolution of pseudogamy. Evolution 39: 294-307.


Moran NA. 1984. The genus Uroleucon in Michigan: key, host records, descriptions of three new species. Journal of the Kansas Entomological Society 57: 596-616.

Moran NA. 1984. Reproductive performance of a specialist herbivore, Uroleucon nigrotibium, on its host and on a non-host. Oikos 42: 171-175.


Moran NA. 1983. Seasonal shifts in host usage in Uroleucon gravicorne (Homoptera: Aphididae) and implications for the evolution of host alternation in aphids. Ecological Entomology 8: 371-382. doi: 10.1111/j.1365-2311.1983.tb00517.x


Moran NA. 1981. Intraspecific variation in herbivore performance and host quality: a field study of Uroleucon caligatum (Homptera: Aphididae) and its Solidago (Asteraceae) hosts. Ecological Entomology 6: 301-306. doi: 10.1111/j.1365-2311.1981.tb00617.x


Hamilton, W. D., P. A. Henderson, and N. A. Moran. 1980. Fluctuations of environment and coevolved antagonist polymorphism as factors in the maintenance of sex. Pages 363-382 in Natural Selection and Social Behavior: recent research and new theory, edited by R. D. Alexander and D. W. Tinkle. Chiron Press, NY.

Moran NA, Hamilton WD. 1980. Low nutritive quality as defense against herbivores. Journal of Theoretical Biology 86: 247-254.


Burley N, Moran NA. 1979. Significance of age and reproductive experience in the mate preferences of feral pigeons, Columba livia. Animal Behaviour 27: 686-698.

Popular Articles

Leonard S and Moran NA. 2021. Engineering bacteria to save honey bees in The Science Breaker: Science Meets Society. 11 FEB 2021 DOI: 10.25250/thescbr.brk471

Moran NA. 2012. Microbial symbiosis and evolution. pages 191-196 in Microbes and Evolution: The World that Darwin Never Saw, edited by R. Kolter and S. Maloy, ASM Press.

Moran NA. 2002. Genome evolution in endosymbiotic bacteria. American Society of Microbiology News 68:499-505.

Moran NA. 2001. Coevolución de bacterias endosimbióticas e insectos. Methode 28: 40-43, Catalan language popular magazine about science (translated).

Moran NA. 2000. The secret to success in sap-sucking insects. Wings (essays on invertebrate conservation) 23: 12-16.

Moran NA. 1992. Quantum leapers: aphids take risks to satisfy ancient tastes. Natural History 4/92: 34-39.