This is a full list of Alvaro’s publications.


Tackling AMR from a multidisciplinary perspective: a primer from education and psychology.
Calvo-Villamañán, A., San Millan Á. & Carrilero. Int Microbiol 2022, doi:

Within-patient evolution of plasmid-mediated antimicrobial resistance.
DelaFuente J, Toribio-Celestino L, Santos-López A, León-Sampedro R, Alonso-del Valle A, Costas C, Hernández-García M, Cui L, Rodriguez-Beltran J, Bikard D, Cantón R, San Millan Á. Nature Ecology & Evolution 2022, doi:

The journey of bacterial genes (News and Views).
San Millán Á. Nature Ecology & Evolution 2022 Mar 28; doi: 10.1038/s41559-022-01713-2

Translational demand is not a major source of plasmid-associated fitness costs.
Rodríguez-Beltrán J, León-Sampedro R, Ramiro-Martínez P, de la Vega C, Baquero F, Levin BR, San Millán Á. Philos Trans R Soc Lond B Biol Sci. 2022 Jan 17;377(1842):20200463. doi: 10.1098/rstb.2020.0463.


Mathematical Models of Plasmid Population Dynamics.
Hernández-Beltrán JCR, San Millán A, Fuentes-Hernández A, Peña-Miller R. Front Microbiol. 2021 Nov 4;12:606396. doi: 10.3389/fmicb.2021.606396.

Staphylococcal phages and pathogenicity islands drive plasmid evolution.
Humphrey S, San Millán Á, Toll-Riera M, Connolly J, Flor-Duro A, Chen J, Ubeda C, MacLean RC, Penadés JR. Nat Commun. 2021 Oct 6;12(1):5845. doi: 10.1038/s41467-021-26101-5.

The bacterial capsule is a gatekeeper for mobile DNA.
Santos-López A, Rodríguez-Beltrán J, San Millán Á. PLoS Biol. 2021 Jul 6;19(7):e3001308. doi: 10.1371/journal.pbio.3001308. eCollection 2021 Jul.

Evolutionary Pathways and Trajectories in Antibiotic Resistance.
Baquero F, Martínez JL, F Lanza V, Rodríguez-Beltrán J, Galán JC, San Millán A, Cantón R, Coque TM. Clin Microbiol Rev. 2021 Jun 30;34(4):e0005019. doi: 10.1128/CMR.00050-19. Online ahead of print.

Variability of plasmid fitness effects contributes to plasmid persistence in bacterial communities.
Alonso-Del Valle A, León-Sampedro R, Rodríguez-Beltrán J, DelaFuente J, Hernández-García M, Ruiz-Garbajosa P, Cantón R, Peña-Miller R, San Millán A. Nat Commun. 2021 May 11;12(1):2653. doi: 10.1038/s41467-021-22849-y.

Pervasive transmission of a carbapenem resistance plasmid in the gut microbiota of hospitalized patients.
León-Sampedro R, DelaFuente J, Díaz-Agero C, Crellen T, Musicha P, Rodríguez-Beltrán J, de la Vega C, Hernández-García M; R-GNOSIS WP5 Study Group, López-Fresneña N, Ruiz-Garbajosa P, Cantón R, Cooper BS, San Millán Á. Nat Microbiol. 2021 May;6(5):606-616. doi: 10.1038/s41564-021-00879-y. Epub 2021 Mar 29.

Collateral sensitivity associated with antibiotic resistance plasmids.
Herencias C, Rodriguez-Beltran J, León-Sampedro R, Alonso-del Valle A, Palkovičová J, Cantón R, San Millán A. eLife. 2021. 10:e65130.

Beyond horizontal gene transfer: the role of plasmids in bacterial evolution.
Rodriguez-Beltran J, DelaFuente JH, León-Sampedro R, MacLean RC & San Millan A. Nature Reviews Microbiology. 2021. DOI: 10.1038/s41579-020-00497-1.


Genetic dominance governs the evolution and spread of mobile genetic elements in bacteria.
Rodríguez-Beltrán J, Sørum V, Toll-Riera M, de la Vega C, Peña-Miller R, San Millán Á. Proc Natl Acad Sci U S A. 2020 Jun 22:202001240. doi: 10.1073/pnas.2001240117.

Quantifying plasmid dynamics using single-cell microfluidics and image bioinformatics.
Hernandez-Beltran JCR, Rodríguez-Beltrán J, San Millán A, Peña-Miller R, Fuentes-Hernández A. Plasmid. 2020 Jun 11:102517. doi: 10.1016/j.plasmid.2020.102517.

Simulating the Influence of Conjugative Plasmids Kinetic Values on the Multilevel Dynamics of Antimicrobial Resistance in a Membrane Computing Model.
Campos M, San Millán Á, Sempere JM, Lanza VF, Coque TM, Llorens C, Baquero F. Antimicrob Agents Chemother. 2020 May 26:AAC.00593-20. doi: 10.1128/AAC.00593-20.



Methods to Study Fitness and Compensatory Adaptation in Plasmid-Carrying Bacteria.
DelaFuente J, Rodriguez-Beltran J, San Millan A. Methods Mol Biol. 020;2075:371-382. doi: 10.1007/978-1-4939-9877-7_26.

The evolution of antibiotic resistance.
MacLean RC, San Millan A. Science. 2019 Sep 13;365(6458):1082-1083. doi: 10.1126/science.aax3879.

Transfer dynamics of Tn6648, a composite integrative conjugative element generated by tandem accretion of Tn5801 and Tn6647 in Enterococcus faecalis.
León-Sampedro R, Fernández-de-Bobadilla MD, San Millán Á, Baquero F, Coque TM. J Antimicrob Chemother. 2019 Jun 21. pii: dkz239. doi: 10.1093/jac/dkz239.



Integrative analysis of fitness and metabolic effects of plasmids in Pseudomonas aeruginosa PAO1.
San Millan A, Toll-Riera M, Qi Q, Betts A, Hopkinson RJ, McCullagh J, MacLean RC. ISME J. 2018 Aug 10. doi: 10.1038/s41396-018-0224-8.

Evolution of Plasmid-Mediated Antibiotic Resistance in the Clinical Context.
San Millan A. Trends Microbiol. 2018 Jul 23. doi: 10.1016/j.tim.2018.06.007.

Testing the Role of Multicopy Plasmids in the Evolution of Antibiotic Resistance.
Escudero JA, MacLean RC, San Millan A. J Vis Exp. 2018 May 2;(135). doi: 10.3791/57386.

Multicopy plasmids allow bacteria to escape from fitness trade-offs during evolutionary innovation.
Rodriguez-Beltran J, Hernandez-Beltran JCR, DelaFuente J, Escudero JA, Fuentes-Hernandez A, MacLean RC, Peña-Miller R, San Millan A. Nature Ecology and Evolution. 2018 Apr 9. doi: 10.1038/s41559-018-0529-z.

Cooperation, competition and antibiotic resistance in bacterial colonies.
Frost I, Smith WPJ, Mitri S, San Millan A, Davit Y, Osborne JM, Pitt-Francis JM, MacLean RC, Foster KR. ISME Journal. 2018 Mar 21. doi: 10.1038/s41396-018-0090-4.

PCR-Based Analysis of ColE1 Plasmids in Clinical Isolates and Metagenomic Samples Reveals Their Importance as Gene Capture Platforms.
Ares-Arroyo M, Bernabe-Balas C, Santos-Lopez A, Baquero MR, Prasad KN, Cid D, Martin-Espada C, San Millan A, Gonzalez-Zorn B. Frontiers in Microbiology. 2018 Mar 16;9:469. doi: 10.3389/fmicb.2018.00469.



Fitness Costs of Plasmids: a Limit to Plasmid Transmission.
San Millan A, MacLean RC. Microbiol Spectr. 2017 Sep;5(5). doi: 10.1128/microbiolspec.MTBP-0016-2017.

A Naturally Occurring Single Nucleotide Polymorphism in a Multicopy Plasmid Produces a Reversible Increase in Antibiotic Resistance.
Santos-Lopez A, Bernabe-Balas C, Ares-Arroyo M, Ortega-Huedo R, Hoefer A, San Millan A, Gonzalez-Zorn B. Antimicrob Agents Chemother. 2017 Jan 24;61(2). pii: e01735-16.



Multicopy plasmids potentiate the evolution of antibiotic resistance in bacteria.
San Millan A, Escudero JA, Gifford DR, Mazel D, MacLean RC. Nature Ecology and Evolution. 2016 Nov 7;1(1):10. doi: 10.1038/s41559-016-0010.
– Selected by Faculty of 1000.
– Highlighted in Nature Microbiology

The Genomic Basis of Evolutionary Innovation in Pseudomonas aeruginosa.
Toll-Riera M, San Millan A, Wagner A, MacLean RC. PLoS Genet. 2016 May 5;12(5):e1006005.



Evaluating the effect of horizontal transmission on the stability of plasmids under different selection regimes.
Peña-Miller R, Rodríguez-González R, MacLean RC, San Millan A. Mobile Genetic Elements. 2015 May 21;5(3):1-5.

Microbial Evolution: Towards Resolving the Plasmid Paradox (Dispatch).
MacLean RC, San Millan A. Current Biology. 2015 Aug 31;25(17):R764-7.

Interactions between horizontally acquired genes create a fitness cost in Pseudomonas aeruginosa.
San Millan A, Toll-Riera M, Qi Q, MacLean RC. Nature Communications. 2015 Apr 21;6:6845.

Small-plasmid-mediated antibiotic resistance is enhanced by increases in plasmid copy number and bacterial fitness.
San Millan A, Santos-Lopez A, Ortega-Huedo R, Bernabe-Balas C, Kennedy SP, Gonzalez-Zorn B.  Antimicrobial Agents and Chemotherapy. 2015;59(6):3335-41.



Positive selection and compensatory adaptation interact to stabilize non-transmissible plasmids.
San Millan A, Peña-Miller R, Toll-Riera M, Halbert ZV, McLean AR, Cooper BS, MacLean RC. Nature Communications. 2014 Oct 10;5:5208.
-Highlighted in Nature Reviews in Microbiology.

Culturable aerobic and facultative bacteria from the gut of the polyphagic dung beetle Thorectes lusitanicus.
Hernández N, Escudero JA, San Millán Á, González-Zorn B, Lobo JM, Verdú JR, Suárez M. Insect Science. 2015 Apr;22(2):178-90.

Positive epistasis between co-infecting plasmids promotes plasmid survival in bacterial populations.
San Millan A, Heilbron K, MacLean RC. ISME Journal. 2014 Mar;8(3):601-12.



SatR is a repressor of fluoroquinolone efflux pump SatAB.
Escudero JA, San Millan A, Montero N, Gutierrez B, Ovejero CM, Carrilero L, González-Zorn B. Antimicrobial Agents and Chemotherapy. 2013 Jul;57(7):3430-3.



Molecular organization of small plasmids bearing blaTEM-1 and conferring resistance to β-lactams in Haemophilus influenzae.
Søndergaard A, San Millan A, Santos-Lopez A, Nielsen SM, Gonzalez-Zorn B, Nørskov-Lauritsen N. Antimicrobial Agents and Chemotherapy. 2012 Sep;56(9):4958-60.

Fitness cost and interference of Arm/Rmt aminoglycoside resistance with the RsmF housekeeping methyltransferases.
Gutierrez B, Escudero JA, San Millan A, Hidalgo L, Carrilero L, Ovejero CM, Santos-Lopez A, Thomas-Lopez D, Gonzalez-Zorn B. Antimicrobial Agents and Chemotherapy. 2012 May;56(5):2335-41.



Fluoroquinolone efflux in Streptococcus suis is mediated by SatAB and not by SmrA.
Escudero JA, San Millan A, Gutierrez B, Hidalgo L, La Ragione RM, AbuOun M, Galimand M, Ferrándiz MJ, Domínguez L, de la Campa AG, Gonzalez-Zorn B. Antimicrobial Agents and Chemotherapy. 2011 Dec;55(12):5850-60.

ArmA methyltransferase in a monophasic Salmonella enterica isolate from food.
Granier SA, Hidalgo L, San Millan A, Escudero JA, Gutierrez B, Brisabois A, Gonzalez-Zorn B. Antimicrobial Agents and Chemotherapy. 2011 Nov;55(11):5262-6.

Plasmid-borne 16S rRNA methylase ArmA in aminoglycoside-resistant Klebsiella pneumoniae in Poland.
Zacharczuk K, Piekarska K, Szych J, Jagielski M, Hidalgo L, San Millán A, Gutiérrez B, Rastawicki W, González-Zorn B, Gierczynski R. Journal of Medical Microbiology. 2011 Sep;60(Pt 9):1306-11

Contribution of ROB-1 and PBP3 mutations to the resistance phenotype of a β-lactamase-positive amoxicillin/clavulanic acid-resistant Haemophilus influenzae carrying plasmid pB1000 in Italy.
San Millan A, Giufré M, Escudero JA, Hidalgo L, Gutierrez B, Cerquetti M, Gonzalez-Zorn B. Journal of Antimicrobial Chemotherapy. 2011 Jan;66(1):96-9.



Haemophilus influenzae clinical isolates with plasmid pB1000 bearing blaROB-1: fitness cost and interspecies dissemination.
San Millan A, Garcia-Cobos S, Escudero JA, Hidalgo L, Gutierrez B, Carrilero L, Campos J, Gonzalez-Zorn B. Antimicrobial Agents and Chemotherapy. 2010 Apr;54(4):1506-11.



Novel genetic environment of qnrB2 associated with TEM-1 and SHV-12 on pB1004, an IncHI2 plasmid, in Salmonella Bredeney BB1047 from Spain.
Gutierrez B, Herrera-Leon S, Escudero JA, Hidalgo L, Gonzalez-Sanz R, Arroyo M, San Millan A, Echeita MA, Gonzalez-Zorn B. Journal of Antimicrobial Chemotherapy. 2009 Dec;64(6):1334-6.

Multiresistance in Pasteurella multocida is mediated by coexistence of small plasmids.
San Millan A, Escudero JA, Gutierrez B, Hidalgo L, Garcia N, Llagostera M, Dominguez L, Gonzalez-Zorn B. Antimicrobial Agents and Chemotherapy. 2009 Aug;53(8):3399-404.

VanB-type Enterococcus faecium clinical isolate successively inducibly resistant to, dependent on, and constitutively resistant to vancomycin.
San Millan A, Depardieu F, Godreuil S, Courvalin P. Antimicrobial Agents and Chemotherapy. 2009 May;53(5):1974-82



Beta-lactam resistance in Haemophilus parasuis is mediated by plasmid pB1000 bearing blaROB-1.
San Millan A, Escudero JA, Catalan A, Nieto S, Farelo F, Gibert M, Moreno MA, Dominguez L, Gonzalez-Zorn B. Antimicrobial Agents and Chemotherapy. 2007 Jun;51(6):2260-4.

First characterization of fluoroquinolone resistance in Streptococcus suis.
Escudero JA, San Millan A, Catalan A, de la Campa AG, Rivero E, Lopez G, Dominguez L, Moreno MA, Gonzalez-Zorn B. Antimicrob Agents Chemother. 2007 Feb;51(2):777-82.