Julin Maloof

image of Julin Maloof



Plant Biology

Offices and Labs

2131 Life Sciences
(530) 752-8077
2115 Life Sciences
(530) 752-8086


1998 PhD Biochemistry University of California, San Francisco
1989 BA Biology Haverford College

Research Interests

How plants sense and respond to their neighbors

Light is essential for plant growth. Perhaps as a consequence, plants have an intricate set of photoreceptors and responses that they use to optimize their development and physiology to suit their light environment. We study the ways in which plants have evolved differences in their light perception and responses that allow them to thrive in different environments. We are interested in both the genetic and molecular basis of variation in light response as well as the adaptive consequences. A combination of molecular and quantitative genetics is used in Arabidopsis, Tomato, and Brassica


2013 AAAS Fellow

Department and Center Affiliations

Department of Plant Biology


Genetics Society of America
American Society of Plant Biologists
American Association for the Advancement of Science

CBS Grad Group Affiliations

Integrated Genetics and Genomics
Plant Biology

Specialties / Focus

Integrated Genetics and Genomics
  • Model Plants
  • Plant Breeding
Plant Biology
  • Cell and Developmental Biology
  • Molecular Biology, Biochemistry, and Genomics


10/7/2014 8:29:18 PM
  • Maloof, J. N., and Kenyon, C. (1998). The Hox gene lin-39 is required during C. elegans vulval induction to select the outcome of Ras signaling. Development 125, 181-190.
  • Maloof, J. N., Whangbo, J., Harris, J. M., Jongeward, G. D., and Kenyon, C. (1999). A Wnt signaling pathway controls hox gene expression and neuroblast migration in C. elegans. Development 126, 37-49.
  • Maloof, J. N., Borevitz, J. O., Weigel, D., and Chory, J. (2000). Natural variation in phytochrome signaling. Semin Cell Dev Biol 11, 523-530.
  • Maloof, JN, JO Borevitz, T Dabi, J Lutes, RB Nehring, JL Redfern, GT Trainer, JM Wilson, T Asami, CC Berry, D Weigel, J Chory. 2001. Natural variation in light sensitivity of Arabidopsis thaliana. Nature Genetics. 29:441-446
  • Friedrichsen, D. M., Nemhauser, J., Muramitsu, T., Maloof, J. N., Alonso, J., Ecker, J. R., Furuya, M., and Chory, J. (2002). Three redundant brassinosteroid early response genes encode putative bHLH transcription factors required for normal growth. Genetics 162, 1445-1456.
  • Borevitz, JO, JN Maloof, J Lutes, T. Davi, JL Redfern, GT Trainer, JD Werner, T Asami, CC Berry, D Weigel, J Chory. 2002. Quantitative trait loci controlling light and hormone response in two accessions of Arabidopsis thaliana. Genetics. 160:683-69
  • Nordborg, M, JO Borevitz, J Bergelson, CC Berry, J Chory, J Hagenblad, M Kreitman, JN Maloof, T Noyes, PJ Oefner, EA Stahl, D Weigel. 2002. The extent of linkage disequilibrium in Arabidopsis thaliana. Nature Genetics. 30:190-193
  • Maloof, J. N. (2003). Genomic approaches to analyzing natural variation in Arabidopsis thaliana. Curr Opin Genet Dev 13, 576-582.
  • Nemhauser, J.L., Maloof, J.N., and Chory, J. (2003). Building integrated models of plant growth and development. Plant Physiol 132, 436-439.
  • Maloof, J. N. (2003). QTL for plant growth and morphology. Curr Opin Plant Biol 6, 85-90.
  • Maloof JN (2004) Plant development: slowing root growth naturally. Curr Biol 14: R395-6.
  • Wolyn DJ, Borevitz JO, Loudet O, Schwartz C, Maloof J, Ecker JR, Berry CC, Chory J (2004). Light-response quantitative trait loci identified with composite interval and eXtreme array mapping in Arabidopsis thaliana. Genetics 167: 907-17.
  • Lu L, Lee YR, Pan R, Maloof JN, & Liu B (2005) An internal motor kinesin is associated with the Golgi apparatus and plays a role in trichome morphogenesis in Arabidopsis. Mol Biol Cell 16: 811-23.
  • Maloof JN (2005) Small but not forgotten. Heredity 96: 1-2.
  • Nozue K, Maloof JN. (2006) Diurnal regulation of plant growth. Plant Cell Environ 29:396-408
  • Balasubramanian S, Sureshkumar S, Agrawal M, Michael TP, Wessinger C, Maloof JN, Clark R, Warthmann N, Chory J, Weigel D (2006). The PHYTOCHROME C photoreceptor gene mediates natural variation in flowering and growth responses of Arabidopsis thaliana. Nat Genet 38: 711-5.
  • Weinig C, Johnston JA, Willis CG, Maloof JN. (2007) Antagonistic multilevel selection on size and architecture in variable density settings. Evolution Int J Org Evolution 61:58-67
  • Nozue K, Covington MF, Duek PD, Lorrain S, Fankhauser C, Harmer SL, Maloof JN. (2007) Rhythmic growth explained by coincidence between internal and external cues. Nature 448:358-61
  • Filiault DL, Wessinger CA, Dinneny JR, Lutes J, Borevitz JO, Weigel D, Chory J, & Maloof JN (2008). Amino acid polymorphisms in Arabidopsis phytochrome B cause differential responses to light. Proc Natl Acad Sci U S A 105: 3157-62.
  • Schwartz C, Balasubramanian S, Warthmann N, Michael TP, Lempe J, Sureshkumar S, Kobayashi Y, Maloof JN, Borevitz JO, Chory J, & Weigel D (2009). Cis-regulatory Changes at FLOWERING LOCUS T Mediate Natural Variation in Flowering Responses of Arabidopsis thaliana. Genetics
  • Jimenez-Gomez JM, & Maloof JN (2009). Sequence diversity in three tomato species: SNPs, markers, and molecular evolution. BMC Plant Biol 9: 85.
  • Jiménez-Gómez JM, & Maloof JN (2009). Plant research accelerates along the (bio)informatics superhighway: symposium on plant sensing, response and adaptation to the environment. EMBO Rep 10: 568-72.
  • Balasubramanian S, Schwartz C, Singh A, Warthmann N, Kim MC, Maloof JN, Loudet O, Trainer GT, Dabi T, Borevitz JO, Chory J, & Weigel D (2009). QTL Mapping in New Arabidopsis thaliana Advanced Intercross-Recombinant Inbred Lines. PLoS ONE 4: e4318.
  • Jiménez-Gómez JM, Wallace AD, and Maloof JN. Network analysis identifies ELF3 as a QTL for the shade avoidance response in Arabidopsis. PLoS Genet 2010 Sep 9; 6(9).
  • Brock MT, Dechaine JM, Iniguez-Luy FL, Maloof JN, Stinchcombe JR, and Weinig C. Floral Genetic Architecture: An Examination of QTL Architecture Underlying Floral (Co)Variation Across Environments. Genetics 2010 Sep 13.
  • Brock MT, Maloof JN, and Weinig C. Genes underlying quantitative variation in ecologically important traits: PIF4 (phytochrome interacting factor 4) is associated with variation in internode length, flowering time, and fruit set in Arabidopsis thaliana. Mol Ecol 2010 Mar; 19(6) 1187-99.
  • Maloof JN. Recent advances in regulation of flowering. F1000 Biol Rep 2010 Feb 11
  • Kerwin, R. E., Jimenez-Gomez, J. M., Fulop, D., Harmer, S. L., Maloof, J. N. & Kliebenstein, D. J. Network Quantitative Trait Loci Mapping of Circadian Clock Outputs Identifies Metabolic Pathway-to-Clock Linkages in Arabidopsis. Plant Cell 23, 471-485 (2011).
  • Holtan, H. E., Bandong, S., Marion, C. M., Adam, L., Tiwari, S., Shen, Y., Maloof, J. N., Maszle, D. R., Ohto, M. A., Preuss, S., Meister, R., Petracek, M., Repetti, P. P., Reuber, T. L., Ratcliffe, O. J. & Khanna, R. BBX32, An Arabidopsis B-box Protein, Functions in Light Signaling by Suppressing HY5-Regulated Gene Expression And Interacting with STH2. Plant Physiol (2011).
  • Chang, C. S., Maloof, J. N. & Wu, S. H. COP1-mediated degradation of BBX22/LZF1 optimizes seedling development in Arabidopsis. Plant Physiol 156, 228-239 (2011).
  • Nozue, K., Harmer, S. L. & Maloof, J. N. Genomic Analysis of Circadian Clock-, Light-, and Growth-Correlated Genes Reveals PHYTOCHROME-INTERACTING FACTOR5 as a Modulator of Auxin Signaling in Arabidopsis. Plant Physiol 156, 357-372 (2011).
  • Stewart, J. L., Maloof, J. N. & Nemhauser, J. L. PIF Genes Mediate the Effect of Sucrose on Seedling Growth Dynamics. PLoS One 6, e19894 (2011).
  • Kim, K., Shin, J., Lee, S. H., Kweon, H. S., Maloof, J. N. & Choi, G. Phytochromes inhibit hypocotyl negative gravitropism by regulating the development of endodermal amyloplasts through phytochrome-interacting factors. Proc Natl Acad Sci U S A 108, 1729-1734 (2011).
  • Arana, M. V., Marín-de la Rosa, N., Maloof, J. N., Blázquez, M. A. & Alabadí, D. Circadian oscillation of gibberellin signaling in Arabidopsis. Proc Natl Acad Sci U S A 108, 9292-9297 (2011).
  • Jiménez-Gómez, J. M., Corwin, J. A., Joseph, B., Maloof, J. N., & Kliebenstein, D. J. (2011). Genomic analysis of QTLs and genes altering natural variation in stochastic noise. PLoS Genetics, 7(9), e1002295. doi:10.1371/journal.pgen.1002295
  • Chitwood, D. H., Headland, L. R., Filiault, D. L., Kumar, R., Jiménez-Gómez, J. M., Schrager, A. V., Park, D. S., et al. (2012). Native Environment Modulates Leaf Size and Response to Simulated Foliar Shade across Wild Tomato Species. PloS one, 7(1), e29570. doi:10.1371/journal.pone.0029570
  • Chitwood, D. H., Headland, L. R., Kumar, R., Peng, J., Maloof, J. N., & Sinha, N. R. (2012). The developmental trajectory of leaflet morphology in wild tomato species. PLANT PHYSIOLOGY. doi:10.1104/pp.111.192518
  • Filiault, D. L., & Maloof, J. N. (2012). A Genome-Wide Association Study Identifies Variants Underlying the Arabidopsis thaliana Shade Avoidance Response. PLoS Genetics, 8(3), e1002589. doi:10.1371/journal.pgen.1002589
  • Seymour, D. K., Filiault, D. L., Henry, I. M., Monson-Miller, J., Ravi, M., Pang, A., Comai, L., et al. (2012). Rapid creation of Arabidopsis doubled haploid lines for quantitative trait locus mapping. Proceedings of the National Academy of Sciences of the United States of America, 109(11), 4227–4232. doi:10.1073/pnas.1117277109
  • Sharlach M, Dahlbeck D, Liu L, Chiu J, Jiménez-Gómez JM, Kimura S, Koenig D, Maloof JN, Sinha N, Minsavage GV, Jones JB, Stall RE, Staskawicz BJ (2012). Fine genetic mapping of RXopJ4, a bacterial spot disease resistance locus from Solanum pennellii LA716. Theor Appl Genet.
  • Palmer CM, Bush SM, Maloof JN. Phenotypic and Developmental Plasticity in Plants. eLS. John Wiley & Sons. Ltd: Chichester; 2012. p. 1–9.
  • Maloof JN, Nozue K, Mumbach MR, Palmer CM (2013). LeafJ: An ImageJ Plugin for Semi-automated Leaf Shape Measurement. JoVE. 2013;(71).
  • Chitwood DH, Maloof JN, Sinha NR (2013). Dynamic transcriptomic profiles between tomato and a wild relative reflect distinct developmental architectures. Plant Physiol. PMCID: PMC3668051
  • Chitwood DH, Kumar R, Headland LR, Ranjan A, Covington MF, Ichihashi Y, Fulop D, Jiménez-Gómez JM, Peng J, Maloof JN, Sinha NR (2013). A quantitative genetic basis for leaf morphology in a set of precisely defined tomato introgression lines. THE PLANT CELL (7):2465–81. PMCID: PMC3753377
  • Koenig D, Jiménez-Gómez JM, Kimura S, Fulop D, Chitwood DH, Headland LR, Kumar R, Covington MF, Devisetty UK, Tat AV, Tohge T, Bolger A, Schneeberger K, Ossowski S, Lanz C, Xiong G, Taylor-Teeples M, Brady SM, Pauly M, Weigel D, Usadel B, Fernie AR, Peng J, Sinha NR, Maloof JN (2013). Comparative transcriptomics reveals patterns of selection in domesticated and wild tomato. Proceedings of the National Academy of Sciences 110(28):E2655–62. PMCID: PMC3710864
  • Chitwood DH, Ranjan A, Martinez CC, Headland LR, Thiem T, Kumar R, Covington MF, Hatcher T, Naylor DT, Zimmerman S, Downs N, Raymundo N, Buckler ES, Maloof JN, Aradhya M, Prins B, Li L, Myles S, Sinha NR (2014). A modern ampelography: a genetic basis for leaf shape and venation patterning in grape. Plant Physiol. 164(1):259–72. PMCID: PMC3875807
  • Ron M, Dorrity MW, de Lucas M, Toal T, Hernandez RI, Little SA, Maloof JN, Klibenstein DJ, Brady SM (2013). Identification of Novel Loci Regulating Inter-specific Variation in Root Morphology and Cellular Development in Tomato. Plant Physiol. PMCID: PMC3668068
  • Chitwood DH, Ranjan A, Kumar R, Ichihashi Y, Zumstein K, Headland LR, Ostria-Gallardo E, Aguilar-Martínez JA, Bush S, Carriedo L, Fulop D, Martinez CC, Peng J, Maloof JN, Sinha NR (2014). Resolving Distinct Genetic Regulators of Tomato Leaf Shape within a Heteroblastic and Ontogenetic Context. THE PLANT CELL.
  • Ichihashi Y, Aguilar-Martínez JA, Farhi M, Chitwood DH, Kumar R, Millon LV, Peng J, Maloof JN, Sinha NR (2014). Evolutionary developmental transcriptomics reveals a gene network module regulating interspecific diversity in plant leaf shape. Proceedings of the National Academy of Sciences. 111(25):E2616–21. PMCID: PMC4078850
  • Devisetty UK, Covington MF, Tat AV, Lekkala S, Maloof JN (2014). Polymorphism Identification and Improved Genome Annotation of Brassica rapa through Deep RNA Sequencing. G3 (Bethesda).
  • Bolger A, Scossa F, Bolger ME, Lanz C, Maumus F, Tohge T, Quesneville H, Alseekh S, Sørensen I, Lichtenstein G, Fich EA, Conte M, Keller H, Schneeberger K, Schwacke R, Ofner I, Vrebalov J, Xu Y, Osorio S, Aflitos SA, Schijlen E, Jiménez-Gómez JM, Ryngajllo M, Kimura S, Kumar R, Koenig D, Headland LR, Maloof JN, Sinha N, van Ham RCHJ, Lankhorst RK, Mao L, Vogel A, Arsova B, Panstruga R, Fei Z, Rose JKC, Zamir D, Carrari F, Giovannoni JJ, Weigel D, Usadel B, Fernie AR. (2014) The genome of the stress-tolerant wild tomato species Solanum pennellii. Nat Genet. 46(9):1034–8.
  • Fitz Gerald JN, Carlson AL, Smith E, Maloof JN, Weigel D, Chory J, Borevitz JO, Swanson RJ (2014). New Arabidopsis advanced intercross recombinant inbred lines reveal female control of nonrandom mating. Plant Physiol. 165(1):175–85. PMCID: PMC4012578


Maloof Lab website


BIS 181 Comparative Genomics (Fall Quarter)
BIS 180L Genome Biology Lab (Spring Quarter)