Majid R. Foolad, Ph.D.

Research

Plant Genetics and Directed Crop Improvement Using Traditional and Contemporary Techniques. Understanding genetic bases of plant response to biotic and abiotic stresses, and genetic controls of fruit nutritional quality. Of particular interest are resistance to diseases early blight and late blight and improved fruit quality in tomato. Significant efforts are devoted to characterization of tomato wild germplasm and introgression of desirable genes for various traits into the cultivated tomato. Conduct genetic mapping, including mapping of QTLs for complex traits, and develop new tomato breeding lines and hybrid cultivars with desirable characteristics using traditional protocols of plant breeding and contemporary techniques of molecular markers and marker-assisted selection. Have released elite inbred lines, experimental hybrids, and/or commercial cultivar of  fresh-market and processing tomatoes.

Teaching

Plant Breeding - HORT 407

Plant Resource Acquisition and Utilization - PLBIO 512

Advanced Plant Breeding - HORT 497B

Independent Undergraduate Research - HORT 296

Independent Undergraduate Research - HORT 496

Professional Background

2003-present  Professor of Plant Genetics, Dept. of Plant Science, Penn State Univ.

2000-2003      Assoc. Professor of Plant Genetics, Dept. of Horticulture, Penn State Univ.

1994-2000      Assist. Professor of Plant Genetics, Dept. of Horticulture, Penn State Univ.

1993-1994      Postdoctoral Associate (Dept. of Pomology), Univ. of California, Davis

1990-1993      Postdoctoral Associate (Dept. of Genetics), Univ. of California, Davis

1986-1990      Research Assistant (Dept. of Vegetable Crops), Univ. of California, Davis

1985-1986      Research Assistant (Dept. of Agron. and Range Sci.), Univ. of California, Davis

Refereed Publications

Jia M and MR Foolad. 2019. Genetic analysis of late blight resistance in a new RIL population of tomato derived from LB-resistant Solanum pimpinellifolium accession PI 270443. Plant Breeding (https:/DOI: 10.1111/pbr.12791)

Paudel S, P Lin, MR Foolad, JG Ali, EG Rajotte, GW Felton. 2019. Induced Plant Defenses Against Herbivory in Cultivated and Wild Tomato. Chemical Ecology 45 (8): 693–707 (https://link.springer.com/article/10.1007/s10886-019-01090-4)

Gao L, I Gonda, H Sun, Q Ma, K Bao, DM Tieman, EA Burzynski-Chang, TL Fish, KA Stromberg, GL Sacks, TW Thannhauser, MR Foolad, MJ Diez, J Blanca, J Canizares, Y Xu, E van der Knaap, S Huang, HJ Klee, JJ Giovannoni, and Z Fei. 2019. The modern tomato genome and a rare allele regulating fruit flavor revealed through pan-genome analysis. Nature Genetics https://doi.org/10.1038/s41588-019-0410-2

Gonda I, H Ashrafi, DA Lyon, SR Strickler, AM Hulse-Kemp, Q Ma, H Sun, K Stoffel, S Futrell, TW Thannhauser, Z Fei, AE Van Deynze, LA Mueller, JJ Giovannoni and MR Foolad. 2019. Sequencing-Based Bin Map Construction of a Tomato Mapping Population, Facilitating High-Resolution Quantitative Trait Loci Detection. The Plant Genome 12(1) 14 pages. http://dx.doi.org/10.3835/plantgenome2018.02.0010

Ohlson EW, H Ashrafi and MR Foolad. 2018. Identification and mapping of late blight resistance QTLs in tomato accession PI 163245. The Plant Genome 11(3), 16 pages. https://doi:10.3835/plantgenome2018.01.0007

Burzynski EA, I Ryona, BI. Reisch, I Gonda, MR Foolad, JJ Giovannoni, GL Sacks. 2018. Quantifying compound × individual matrix effects in HS-SPME-GC-MS of plant populations. Molecules 23: 2436 (12 pages); https://doi.org/10.3390/molecules23102436

Sullenberger MT and MR Foolad. 2018. Genetic characterization of late blight resistance in Solanum pimpinellifolium accession PI 270442. Adv Studies Biol 10: 13-32. https://doi.org/10.12988/asb.2018.71231

Sullenberger, MT, M Jia, S Gao and MR Foolad. 2018. Genetics analysis of late blight resistance in Solanum pimpinellifolium accession PI 270441: heritability and response to selection. Plant Breeding 137:89–96. https://doi.org/10.1111/pbr.12561

Ohlson, EW and MR Foolad. 2016. Genetic analysis of resistance to tomato late blight in Solanum pimpinellifolium accession PI 163245. Plant Breeding 135: 391-398. doi: 10.1111/pbr.12366

Foolad, MR and L. Zhang. 2015. A recombinant inbred line population of tomato and its genetic map constructed based on a Solanum lycopersicum ´ S. pimpinellifolium cross. Adv Studies Biol 7: 441 – 471.

Ohlson, EW and Foolad, MR. 2015. Heritability of late blight resistance conferred by Solanum pimpinellifolium accession PI 224710. Plant Breeding 134: 461–467.

Foolad, MR, M. Sullenberger and H. Ashrafi. 2015. Detached-leaflets evaluation of tomato germplasm for late blight resistance and its correspondence with field and greenhouse screenings. Plant Disease 99:718-722.

Ashrafi, H. and Foolad, MR. 2015. Characterization of early blight resistance in a recombinant inbred line population of tomato: II. Identification of QTLs and their co-localization with candidate resistance genes. Adv Studies Biol 7: 149-168.

Foolad, MR and H. Ashrafi. 2015. Characterization of early blight resistance in a recombinant inbred line population of tomato: I. Heritability and Trait Correlations. Adv Studies Biol 7: 131-148.

Foolad, MR, M. Sullenberger, E. Ohlson and BC Gugino. 2014. Response of Accessions within Tomato Wild Species, Solanum pimpinellifolium to Late Blight. Plant Breeding 133:401-411; doi:10.1111/pbr.12172

Kinkade MP and MR Foolad. 2013. Validation and fine mapping of lyc12.1, a QTL for increaed tomato fruit lycopene content. Theor Appl Genet DOI: 10.1007/s00122-013-2126-5

Ashraf M and MR Foolad. 2013. Crop breeding for salt tolerance in the era of molecular markers and marker-assisted selection. Plant Breeding 132, 10–20.

 Merk HL, H Ashrafi and MR Foolad. 2012. Selective genotyping to identify late blight resistance genes in an accession of the tomato wild species Solanum pimpinellifolium. Euphytica: DOI 10.1007/s10681-012-0729-6

Ashrafi H*, MP Kinkade*, H Merk* and MR Foolad.  2012. Identification of novel QTLs for increased lycopene content and other fruit quality traits in a tomato RIL population. Molecular Breeding: 30:549–567 DOI: 10.1007/s11032-011-9643-1

Pirasteh Anosheh, H, Y Emam, M. Ashraf  and MR Foolad. 2012. Exogenous application of salicylic acid and chlormequat chloride alleviates negative effects of drought stress in wheat. Adv Studies in Biology 4:501 - 520

Nowicki M, MR Foolad, M Nowakowska and EU Kozik. 2012. Potato and tomato late blight caused by Phytophthora infestans: An overview of pathology and resistance breeding. Plant Disease 96: 4-17.

 Foolad MR and DR Panthee. 2012. Marker-assisted selection in tomato breeding. Cri. Rev. Plant Sci. 31: 93-123.

 Merk HL* and MR Foolad. 2011. Parent-offspring correlation estimate of heritability for late blight resistance conferred by an accession of the tomato wild species Solanum pimpinellifolium. Plant Breeding: 131: 203-210.

Jamil A, M Ashraf and MR Foolad.  2011. Gene expression profiling of plants under salt stress. Cri. Rev. Plant Sci. 30: 30:435–458

 Ashraf M, NA Akram, F Al-Qurainy and MR Foolad.  2011. Drought tolerance: Roles of organic osmolytes, plant growth regulators and mineral nutrients. Adv. Agron. 111: 249-296.

Ashraf M, NA Akram*, RN Arteca and MR Foolad.  2010. The physiological, biochemical and molecular roles of brassinosteroids and salicylic acid in plant processes and salt tolerance. Cri. Rev. Plant Sci. 29:1-29.

 Ashrafi H, MP Kinkade and MR Foolad.  2009. A new genetic linkage map of tomato based on a Solanum lycopersicum × S. pimpinellifolium RIL population displaying locations of candidate resistance ESTs. Genome 52: 935-956.

 Sharma A, L Zhang, D Niño-Liu, H Ashrafi and MR Foolad.  2008.  A Solanum lycopersicum ´ S. pimpinellifolium linkage map of tomato displaying genomic locations of R-genes, RGAs and candidate resistance/defense-response ESTs. Int'l. J. Plant Genomics, vol. 2008, Article ID 926090, 18 pages, 2008. doi:10.1155/2008/926090.

 Foolad MR, H Merk and H Ashrafi.  2008.  Genetics, genomics and breeding of late blight and early blight resistance in tomato. Cri. Rev. Plant Sci. 27: 75-107

 Foolad MR, L. Zhang  and P Subbiah. 2007.  Common QTLs affect the rate of tomato seed germination under different stress and non-stress conditions. Int'l. J. Plant Genomics, vol. 2007, Article ID 97386, 10 pages, 2007. doi:10.1155/2007/97386.

 Foolad MR. 2007. Genome mapping and molecular breeding of tomato. International Journal of Plant Genomics, vol. 2007, Article ID 64358, 52 pages. doi:10.1155/2007/64358

Wahid A, S Gelani, M Ashraf and MR Foolad.  2007.  Heat tolerance in plants: A critical review. Env. Exp. Bot. 6: 199–223

 Ashraf M and MR Foolad. 2007. Roles of glycine betaine and proline in improving plant abiotic stress resistance. Environ. Exp. Bot. 59:206-216.

 Foolad MR and A Sharma. 2005. Molecular markers as selection tools in tomato breeding. Acta Hort 695: 225-240.

 Foolad MR, A Sharma, H Ashrafi, and GY Lin. 2005. Genetics of early blight resistance in tomato. Acta Hort 695: 397-406.

 Foolad MR. 2005. Recent development in stress tolerance breeding in tomato. In: Abiotic Stresses: Plant Resistance through Breeding and Molecular Approaches. M Ashraf and PJC Harris, eds. The Haworth Press Inc., New York, USA. pp. 613-684.

 Ashraf M and MR Foolad. 2005. Pre-sowing seed treatment – a shotgun approach to improve germination, plant growth and crop yield under saline and non-saline conditions. Adv. Agron. 225-271.

 Foolad MR. 2004. Recent advances in genetics of salt tolerance in tomato. Plant Cell, Tissue and Organ Cultur: Reviews of Plant Biotechnology and Applied Genetics 76: 101-119.

 Hyman JR, J Gaus, and MR Foolad. 2004. A rapid and accurate method for estimating tomato lycopene content by measuring chromaticity values of fruit puree. JASHS 129: 717-723.

 Zhang LP, GY Lin, and MR Foolad. 2003. QTL comparison of salt tolerance during seed germination and vegetative growth in a Lycopersicon esculentum ´ L. pimpinellifolium RIL population. Acta Hort. 618: 59-67.

 Foolad MR, L Zhang, and P Subbiah. 2003. Relationships among cold, salt and drought tolerance during seed germination in tomato: Inheritance and QTL mapping. Acta Hort. 618: 47-57.

 Zhang L, GY Lin, D Niño-Liu, MR Foolad. 2003. Mapping QTLs conferring early blight (Alternaria solani) resistance in a Lycopersicon esculentum ´ L. hirsutum cross by selective genotyping. Molec. Breeding 12: 3-19.

 Foolad MR, L Zhang, P. Subbiah. 2003. Genetics of drought tolerance during seed germination in tomato: Inheritance and QTL mapping. Genome 46: 536-545.

 Niño-Liu D, LP Zhang, and MR Foolad. 2003. Sequence Comparison and Characterization of Mapped resistance gene analogs Markers in Tomato. Acta Hort. 625:49-58.

 Foolad MR, P Subbiah, C Kramer, G Hargrave, LP Zhang, GY Lin. 2003. Genetic relationships between cold, salt and drought tolerance during seed germination in an interspecific cross of tomato. Euphytica 130: 199-206.

 Bliss FA, S Arulsekar, MR Foolad, V Becerra, C Thormann, AM Gillen, ML Warburton, AM Dandekar, GM Kocsisne, KK Mydin. 2002. An expanded genetic linkage map of Prunus based on an interspecific cross between almond and peach. Genome 45: 520-529.

 Foolad MR, P Subbiah, GS Ghangas. 2002. Parent-offspring correlation estimate of heritability for early blight resistance in tomato, Lycopersicon esculentum Mill. Euphytica 126: 291-297.

 Foolad MR, L Zhang, A. Khan, D Niño-Liu, GY Lin. 2002. Identification of QTLs for early blight (Alternaria solani) resistance in tomato using backcross populations of a Lycopersicon esculentum ´ L. hirsutum cross. Theor. Appl. Genet. 104: 945-958.

 Zhang L, A Khan, D Niño-Liu, MR Foolad 2002. A molecular linkage map of tomato displaying chromosomal locations of resistance gene analogs identified based on a Lycopersicon esculentum ´ L. hirsutum cross. Genome 45: 133-146.

 Foolad MR, GY Lin. 2001. Genetic analysis of cold tolerance during vegetative growth in tomato, Lycopersicon esculentum Mill. Euphytica 122: 105-111.

 Foolad MR, L Zhang, GY Lin. 2001. Identification and validation of qtls for salt tolerance during vegetative growth in tomato by selective genotyping. Genome 44: 444-454.

 Foolad, MR and GY Lin. 2001. Relationship between cold tolerance during seed germination and vegetative growth in tomato. Analysis of response and correlated response to selection. J. Amer. Soc. Hort. Sci. 126: 216-220.

Foolad, MR and GY Lin. 2001. Heritability of early blight resistance in a Lycopersicon esculentum ´ L. hirsutum cross estimated by correlation between parent and progeny. Plant Breeding 120: 173-177.

 Foolad MR 2000. Genetics bases of salt tolerance and cold tolerance in tomato. Current Topics in Plant Biology 2: 35-49.

Foolad MR, GY Lin. 2000. Relationship between cold tolerance during seed germination and vegetative growth in tomato. Germplasm evaluation. J. Amer. Soc. Hort. Sci. 125: 679-683.

 Foolad MR, N Ntahimpera, BJ Christ, GY Lin. 2000. Comparison between field, greenhouse, and detached-leaflet evaluations of tomato germplasm for early blight resistance. Plant Disease 84: 967-972.

 Foolad MR, Chen FQ, Lin GY 1999. RFLP mapping of qtls conferring salt tolerance during vegetative growth in an interspecific cross of tomato. Theor. Appl. Genet. 99: 235-243.

 Foolad MR 1999. Comparison of salt tolerance during seed germination and vegetative growth in tomato by QTL mapping. Genome 42: 727-734.

 Foolad MR, Lin GY, Chen FQ 1999. Comparison of QTLs for seed germination under non-stress, cold stress and salt stress in tomato. Plant Breeding 118: 167-173.

 Foolad MR, Lin GY 1999. Relationships between cold- and salt-tolerance during seed germination in tomato: Germplasm evaluation. Plant Breeding 118: 45-48.

 Foolad MR, Hyman JR, Lin GY 1999. Relationships between cold- and salt-tolerance during seed germination in tomato: Analysis of response and correlated response to selection. Plant Breeding 118: 49-52.

 Foolad MR 1999. Genetics of salt tolerance and cold tolerance in tomato: Quantitative analysis and qtl mapping. Plant Biotechnology 16: 55-64.

 Chen FQ, Foolad MR, Hyman JR, St. Clair D, Beelman R. 1999. Mapping of qtls for lycopene and other fruit traits in a Lycopersicon esculentum ´ L. pimpinellifolium cross and comparison of QTLs across tomato species. Molec. Breeding 5: 283-299.

 Chen FQ, Foolad MR 1999. Nucellar cell-specific expression of a lipid transfer protein (LTP) gene in barley (Hordeum vulgare L.). Plant Cell Rprts. 18: 445-450.

 Chen FQ, Foolad MR 1999. A molecular linkage map of tomato based on a cross between Lycopersicon esculentum and L. pimpinellifolium and its comparison with other molecular maps of tomato. Genome 42: 94-103.

 Foolad MR, Chen FQ, Lin GY 1998. RFLP mapping of qtls conferring cold tolerance during germination in an interspecific cross of tomato. Molec. Breed. 4:519-529

 Foolad MR, Chen FQ, Lin GY 1998. RFLP mapping of qtls conferring salt tolerance during germination in an interspecific cross of tomato. Theor. Appl. Genet. 97:1133-1144

 Foolad MR, Lin GY 1998. Genetic analysis of low temperature tolerance during germination in tomato, Lycopersicon esculentum Mill. Plant Breeding 117:171-176

 Foolad MR, Chen FQ 1998. RAPD markers associated with salt tolerance in an interspecific cross of tomato (Lycopersicon esculentum ´ L. pennellii). Plant Cell Rprts 17:306-312

Chen FQ, MR Foolad. 1997. Molecular organization of a gene in barley which encodes a protein similar to aspartic protease and its specific expression in nucellar cells during degeneration. Plant Molec. Biol. 35:821-831

 Foolad MR, Lin GY 1997 Absence of a relationship between salt tolerance during germination and vegetative growth in tomato, Lycopersicon esculentum Mill. Plant Breeding 116:363-367

 Foolad MR, Stoltz T, Dervinis C, Rodriguez RL, Jones RA 1997.  Mapping QTLs conferring salt tolerance during germination in tomato by selective genotyping.  Molecular Breeding 3:269-277

 Foolad MR, Lin GY 1997 Genetic potential for salt tolerance during germination in lycopersicon species. HortScience 32:296-300

 Foolad MR 1997 Genetic basis of physiological traits related to salt tolerance in tomato, Lycopersicon esculentum Mill. Plant Breeding 116:53-58

 Foolad MR 1996 Response to selection for salt tolerance during germination in tomato seed derived from P.I. 174263.  J. Am. Soc. Hort. Sci. 121:1006-1011

 Foolad MR 1996 Genetic analysis of salt tolerance during vegetative growth in tomato, Lycopersicon esculentum Mill.  Plant Breeding 115:245-250

 Foolad MR 1996 Unilateral incompatibility as a major cause of skewed segregation in an interspecific cross of tomato. Plant Cell Reports 15:627-633

 Foolad MR, Arulsekar S, Becerra V, Bliss FA 1995  A genetic map of Prunus based on an interspecific cross between peach and almond Theor. Appl. Genet. 91: 262-269

 Hittalmani S, Foolad MR, Mew T, Rodriguez RL, Huang N 1995  Development of a PCR-based marker to identify rice blast resistance gene, pi-2(t), in a segregating population. Theor. Appl. Genet. 91: 9-14

 Foolad MR and Jones RA 1993  Mapping salt-tolerance genes in tomato (Lycopersicon esculentum) using trait-based marker analysis. Theor. Appl. Genet. 87:184-192

 Karrer EE, Chandler JC, Foolad MR and Rodriguez RL 1993  Correlation between a-amylase gene expression and seedling vigor in rice. Euphytica  66: 163-169

 Foolad MR , Jones RA, and Rodriguez RL 1993   RAPD markers for constructing intraspecific tomato genetic maps. Plant Cell Reports 12: 293-297

 Kishimoto M, Foolad MR, Shimosaka E, Matsuura S. and Saito A 1993  Alignment of molecular and classical linkage maps of rice, Oryza sativa L. Plant Cell Reports  12: 457-461

 Foolad MR and Jones RA 1992  Parent-offspring regression estimates of heritability for salt tolerance during germination in tomato. Crop Sci. 32: 439-442

 Foolad MR and Jones RA 1992  Models to estimate maternally controlled genetic variation in quantitative seed traits. Theor. Appl. Genet. 83: 360-366

 Ranjahn S, Litts JC, Foolad MR and Rodriguez RL 1991  Chromosomal localization and genomic organization of a-amylase genes in rice (Oryza sativa L.). Theor. Appl. Genet. 82: 481-488

 Foolad MR and Jones RA 1991  Genetic analysis of salt tolerance during germination in Lycopersicon.  Theor. Appl. Genet. 81: 321-326

 Kozik E, Foolad MR and Jones RA 1991  Genetic analysis of resistance to Phytophthora root rot in tomato (Lycopersicon esculentum Mill).  Plant Breeding 106: 27-32