Research Focus: We study how roots and soil fungi interact with soils and whole-plant processes to create functional plant strategies in complex ecosystems.
Key Words: Fine roots, mycorrhizal fungi, ecosystem ecology, global change biology, forest ecology, plant physiology, urban forestry, ecological modelling, biogeochemistry
Key Words: Fine roots, mycorrhizal fungi, ecosystem ecology, global change biology, forest ecology, plant physiology, urban forestry, ecological modelling, biogeochemistry
Evolutionary history and global variation in fine-root traits

Working with a collaborative group across Europe, China and the North America, this project utilizes an extensive collection of woody plant species at The Morton Arboretum, with notably rich coverage of oak species (genus Quercus) and a diverse collection of gymnosperm species that are currently underrepresented in most global surveys. With this foundation, we are addressing basic research questions regarding how the diverse fine-root strategies observed in plants today have evolved across species, ecosystems, and biomes. We are further interested in how these different strategies impact plant performance, ecosystem nutrient cycling and soil carbon storage. Our recent work has suggested a reduction in plant reliance on symbiotic soil fungi (i.e. mycorrhizal fungi) among more recently derived angiosperm species and these changes within the angiosperm lineage have led to broad shifts in plant strategies for belowground resource acquisition. However, it is unclear if this pattern is consistent across all plants and if the pattern of narrowing fine-root diameter and reduced fungal reliance found in angiosperms is the same with different types of mycorrhizal associations. We also have precious little understanding of how fine-root traits vary within a particular genus or species along environmental gradients. At present, exceptions to the rule seem nearly as common as the expected pattern itself suggesting that there is much left to be understood and many questions left to be answered.
Figure (McCormack and Kaproth et al. unpublished). Phylogenetic tree along with summary information of fine-root trait variation for individual root orders (1st, 2nd 3rd order) among 20 species of oak (Quercus). Oak phylogeny was derived from Hipp et al. (2018) and the names of the major oak sections included are given at their respective branches. Range of variation is displayed for each fine-root trait with relatively larger values indicated in black and increasingly smaller values indicated progressively from grey to white. The coefficient of variation calculated across the 20 species is provided at the bottom of each trait column. Leaf habit of each species are also indicated with deciduous, brevi-deciduous, and evergreen indicated in light, medium, and dark green.
Relevant Publications
McCormack ML*, MA Kaproth*, E Carlson, J Cavender-Bares, PG Kennedy. Climate and phylogenetic history structure morphological and architectural trait variation among fine-root orders. New Phytologist, (2020).
Guerrero-Ramirez N, Mommer L, Freschet GT, Iversen CM, McCormack ML, Kattge J, Poorter H, van der Plas F, Bergmann J, Kuyper TW, ... A Weigelt. Global Root Traits (GRooT) Database. Global Ecology and Biogeography (2020).
Bergmann J, A Weigelt, F van der Plas, DC Laughlin, TW Kuyper, N Guerrero-Rameriz, OJ Valverde-Barrantes, H Bruelheide, GT Freschet, CM Iversen, J Kattge, ML McCormack, IC Meier, MC Rillig, C Roumet, M Semchenko, CJ Sweeney, J van Ruijven, LM York, L Mommer, The fungal collaboration gradient dominates the root economics space in plants. Science Advances, 6: eaba3756 (2020).
Defrenne CE, ML McCormack, SD Addo-Danso, SW Simard. Intraspecific fine-root trait-environment relationships across interior Douglas-fir forests of western Canada. Plants, 8: 199 (2019).
See CR, ML McCormack, SE Hobbie, H Flores-Moreno, WL Silver, PG Kennedy. Global patterns in fine-root decomposition.... Ecology Letters, 22: 946-953 (2019).
Li F, H Hu, ML McCormack, D Feng, X Liu, W Bao. A community-level economics spectrum of fine-roots driven by nutrient limitations in subalpine forests. Journal of Ecology, 107: 1238-1249 (2019).
Li L, ML McCormack, F Chen, H Wang, Z Ma, D Guo. Different responses of absorptive roots and arbuscular mycorrhizal fungi.... Forest Ecology and Management, 433: 64-7 (2019).
Caplan JS, SJ Meiners, H Flores-Moreno, ML McCormack. Fine-root traits are linked to species dynamics in a successional plant community. Ecology, 100: e02588 (2019).
Wang C, ML McCormack, D Guo, J Li. Angiosperms and gymnosperms differ in patterns of root carbon allocation along environmental gradients. Journal of Biogeography, 46: 123-133 (2019).
Mao Z, Y Wang, ML McCormack , N Rowe, X Deng, X Yang, S Xia, J Nespoulous, RC Sidle, D Guo, A Stokes. Mechanical properties of fine roots.... Annals of Botany, mcy076 (2018).
Ma Z, D Guo, X Xu, M Lu, RD Bardgett, DM Eissenstat, ML McCormack, LO Hedin. Evolutionary history resolves global organization of root functional traits. Nature, 555: 94-97 (2018).
Li H, B Liu, ML McCormack, B Zhu, Z Ma, D Guo. Diverse belowground resource strategies underlie plant species coexistence and spatial distribution. New Phytologist, 216: 1140-1150 (2017).
McCormack ML, D Guo, CM Iversen, W Chen, DM Eissenstat, CW Fernandez, L Li, C Ma, Z Ma, H Poorter, PB Reich, M Zadworny, AE Zanne. Building a better foundation.... New Phytologist, 215: 27-37 (2017).
Freschet GT, OJ Valverde-Barrantes, CM Tucker, JM Craine, ML McCormack et al..... Climate, soil and plant functional types as drivers of global fine-root trait variation. Journal of Ecology, 105: 1182-1196 (2017).
Figure (McCormack and Kaproth et al. unpublished). Phylogenetic tree along with summary information of fine-root trait variation for individual root orders (1st, 2nd 3rd order) among 20 species of oak (Quercus). Oak phylogeny was derived from Hipp et al. (2018) and the names of the major oak sections included are given at their respective branches. Range of variation is displayed for each fine-root trait with relatively larger values indicated in black and increasingly smaller values indicated progressively from grey to white. The coefficient of variation calculated across the 20 species is provided at the bottom of each trait column. Leaf habit of each species are also indicated with deciduous, brevi-deciduous, and evergreen indicated in light, medium, and dark green.
Relevant Publications
McCormack ML*, MA Kaproth*, E Carlson, J Cavender-Bares, PG Kennedy. Climate and phylogenetic history structure morphological and architectural trait variation among fine-root orders. New Phytologist, (2020).
Guerrero-Ramirez N, Mommer L, Freschet GT, Iversen CM, McCormack ML, Kattge J, Poorter H, van der Plas F, Bergmann J, Kuyper TW, ... A Weigelt. Global Root Traits (GRooT) Database. Global Ecology and Biogeography (2020).
Bergmann J, A Weigelt, F van der Plas, DC Laughlin, TW Kuyper, N Guerrero-Rameriz, OJ Valverde-Barrantes, H Bruelheide, GT Freschet, CM Iversen, J Kattge, ML McCormack, IC Meier, MC Rillig, C Roumet, M Semchenko, CJ Sweeney, J van Ruijven, LM York, L Mommer, The fungal collaboration gradient dominates the root economics space in plants. Science Advances, 6: eaba3756 (2020).
Defrenne CE, ML McCormack, SD Addo-Danso, SW Simard. Intraspecific fine-root trait-environment relationships across interior Douglas-fir forests of western Canada. Plants, 8: 199 (2019).
See CR, ML McCormack, SE Hobbie, H Flores-Moreno, WL Silver, PG Kennedy. Global patterns in fine-root decomposition.... Ecology Letters, 22: 946-953 (2019).
Li F, H Hu, ML McCormack, D Feng, X Liu, W Bao. A community-level economics spectrum of fine-roots driven by nutrient limitations in subalpine forests. Journal of Ecology, 107: 1238-1249 (2019).
Li L, ML McCormack, F Chen, H Wang, Z Ma, D Guo. Different responses of absorptive roots and arbuscular mycorrhizal fungi.... Forest Ecology and Management, 433: 64-7 (2019).
Caplan JS, SJ Meiners, H Flores-Moreno, ML McCormack. Fine-root traits are linked to species dynamics in a successional plant community. Ecology, 100: e02588 (2019).
Wang C, ML McCormack, D Guo, J Li. Angiosperms and gymnosperms differ in patterns of root carbon allocation along environmental gradients. Journal of Biogeography, 46: 123-133 (2019).
Mao Z, Y Wang, ML McCormack , N Rowe, X Deng, X Yang, S Xia, J Nespoulous, RC Sidle, D Guo, A Stokes. Mechanical properties of fine roots.... Annals of Botany, mcy076 (2018).
Ma Z, D Guo, X Xu, M Lu, RD Bardgett, DM Eissenstat, ML McCormack, LO Hedin. Evolutionary history resolves global organization of root functional traits. Nature, 555: 94-97 (2018).
Li H, B Liu, ML McCormack, B Zhu, Z Ma, D Guo. Diverse belowground resource strategies underlie plant species coexistence and spatial distribution. New Phytologist, 216: 1140-1150 (2017).
McCormack ML, D Guo, CM Iversen, W Chen, DM Eissenstat, CW Fernandez, L Li, C Ma, Z Ma, H Poorter, PB Reich, M Zadworny, AE Zanne. Building a better foundation.... New Phytologist, 215: 27-37 (2017).
Freschet GT, OJ Valverde-Barrantes, CM Tucker, JM Craine, ML McCormack et al..... Climate, soil and plant functional types as drivers of global fine-root trait variation. Journal of Ecology, 105: 1182-1196 (2017).
Fine-root dynamics and aboveground-belowground linkages in trees

Figure. The importance and seasonal patterns of leaf emergence, maturation, and senescence are well know for most tree species (spring, summer, fall, winter seasons shown in top panels), but there is relatively little information concerning the patterns and drivers of belowground processes such as root growth and fungal dynamics (bottom panels) in most terrestrial ecosystems.
Together with the leaves and stems that we more commonly observe aboveground, the production and turnover of roots belowground play important role mediating carbon and nutrient cycling within plants and across terrestrial ecosystems. However, basic patterns of phenology, lifespan, and decomposition of fine roots are poorly understood. In fact, compared to leaves aboveground, we know quite little about how different plant species allocate resources to build and maintain roots belowground, how temporal patterns in resource allocation to roots are linked with other components of trees and forests, and how these differences manifest diverse strategies for soil resource acquisition across seasons and years. Through long-term observations established in mature forestry plots at The Morton Arboretum as well as through external collaborations, we are working to identify and understand linkages between fine-root dynamics belowground and broader patterns of plant growth and resource investment belowground. Moreover, we are targeting our observations across a diverse suite of species representing different taxonomic groups, mycorrhizal associations, woody anatomy, fine-root traits, and whole-plant traits. From this we can develop broader understanding of functional plant variation across major plant groups that are especially relevant for the development of forest ecosystem models and terrestrial biosphere models.
Relevant Publications
Mueller KE, DR LeCain, ML McCormack, DM Blumenthal, M Carlson, E Pendall. Root responses to climate change in a semiarid grassland. Journal of Ecology, Journal of Ecology, 106: 2176-2189 (2018).
McCormack ML, CW Fernandez, H Brooks, SG Pritchard. Production dynamics of Cenococcum geophilum ectomycorrhizas .... Fungal Ecology, 26: 11-19 (2017).
Kou L, ML McCormack, W Chen, D Guo, H Wang, W Gao, H Yang, S Li. Ectomycorrhizal morphotype, nitrogen and root distribution mediate lifespan of absorptive roots. Plant and Soil, 411: 261 (2017).
McCormack ML, KP Gaines, MP Pastore, DM Eissenstat. Early season root production in relation to leaf production among six diverse temperate tree species. Plant and Soil, 389: 121-129 (2015).
McCormack ML, TS Adams, EAH Smithwick, DM Eissenstat. Variability in root production and root turnover alter estimates of fine root turnover rate. Ecology 95: 2224-2235 (2014).
Functional trade-offs in the root-mycorrhizal symbiosis

The function of fine roots belowground is intricately linked with mycorrhizal fungi reflecting a symbiosis that has existed since shortly after plants emerged on land. With the exchange of carbohydrates from the plant and belowground resources from the fungi, both partners may realize significant benefit. However, the trade-offs and potential advantages of greater or lesser reliance by plants on mycorrhizal fungi are not well understood. At the same time, these trade-off largely determine competitive plant strategies for belowground resource acquisition strategies across environments. Our ongoing work aims to quantify functional trade-offs between fine roots and their mycorrhizal partners recognizing that the relative benefit to each partner likely changes with environmental conditions and based on the individual plant and fungal species involved.
Figure. Root tips of pine colonized by the ectomycorrhizal fugus Tuber sp. Note the fungal hyphae emanating from the white tips.
Relevant Publications
Bergmann J, A Weigelt, F van der Plas, DC Laughlin, TW Kuyper, N Guerrero-Rameriz, OJ Valverde-Barrantes, H Bruelheide, GT Freschet, CM Iversen, J Kattge, ML McCormack, IC Meier, MC Rillig, C Roumet, M Semchenko, CJ Sweeney, J van Ruijven, LM York, L Mommer, The fungal collaboration gradient dominates the root economics space in plants. Science Advances, 6: eaba3756 (2020).
McCormack ML and CM Iversen. Physical and functional constraints on viable belowground acquisition strategies. Frontiers in Plant Science, 10: 1-12 (2019).
Li L, ML McCormack, F Chen, H Wang, Z Ma, D Guo. Different responses of absorptive roots and arbuscular mycorrhizal fungi.... Forest Ecology and Management, 433: 64-7 (2019).
Zhu K and ML McCormack, RA Lankau, JF Egan, N Wurzburger. Ectomycorrhizal trees with high carbon‐to‐nitrogen soils driven by smaller nitrogen not larger carbon stocks. Journal of Ecology, 106: 524-525 (2018).
Lin G, ML McCormack, C Ma, D Guo. Similar soil carbon but contrasting modes of nitrogen cycling between arbuscular mycorrhizal and ectomycorrhizal forests. New Phytologist, 213: 1440-1451 (2017).
Li H, B Liu, ML McCormack, B Zhu, Z Ma, D Guo. Diverse belowground resource strategies underlie plant species coexistence and spatial distribution. New Phytologist, 216: 1140-1150 (2017).
Cheng L, DM Eissenstat, R Koide, J DeForest, ML McCormack, T Adams, W Chen, X Wei, L Li. Mycorrhizal fungi and roots are complementary in foraging within nutrient patches. Ecology, 97: 2815-2823 (2016).
Figure. Root tips of pine colonized by the ectomycorrhizal fugus Tuber sp. Note the fungal hyphae emanating from the white tips.
Relevant Publications
Bergmann J, A Weigelt, F van der Plas, DC Laughlin, TW Kuyper, N Guerrero-Rameriz, OJ Valverde-Barrantes, H Bruelheide, GT Freschet, CM Iversen, J Kattge, ML McCormack, IC Meier, MC Rillig, C Roumet, M Semchenko, CJ Sweeney, J van Ruijven, LM York, L Mommer, The fungal collaboration gradient dominates the root economics space in plants. Science Advances, 6: eaba3756 (2020).
McCormack ML and CM Iversen. Physical and functional constraints on viable belowground acquisition strategies. Frontiers in Plant Science, 10: 1-12 (2019).
Li L, ML McCormack, F Chen, H Wang, Z Ma, D Guo. Different responses of absorptive roots and arbuscular mycorrhizal fungi.... Forest Ecology and Management, 433: 64-7 (2019).
Zhu K and ML McCormack, RA Lankau, JF Egan, N Wurzburger. Ectomycorrhizal trees with high carbon‐to‐nitrogen soils driven by smaller nitrogen not larger carbon stocks. Journal of Ecology, 106: 524-525 (2018).
Lin G, ML McCormack, C Ma, D Guo. Similar soil carbon but contrasting modes of nitrogen cycling between arbuscular mycorrhizal and ectomycorrhizal forests. New Phytologist, 213: 1440-1451 (2017).
Li H, B Liu, ML McCormack, B Zhu, Z Ma, D Guo. Diverse belowground resource strategies underlie plant species coexistence and spatial distribution. New Phytologist, 216: 1140-1150 (2017).
Cheng L, DM Eissenstat, R Koide, J DeForest, ML McCormack, T Adams, W Chen, X Wei, L Li. Mycorrhizal fungi and roots are complementary in foraging within nutrient patches. Ecology, 97: 2815-2823 (2016).
Trees and water stress

Too little and too much water both present challenges to trees. As a result, trees have developed many strategies to adapt to drought and waterlogging conditions. Aspects of physiological, anatomical, and morphological adapations are well studied aboveground. Despite being the primary point of contact between plants and soil water, we have comparatively poor understanding of how roots respond to water stress and and how unique adaptations at the root level can facilite better tree growth under stressful conditions.
We are interested to address this knowledge gap and better understand how roots mediate plant responses to drought and waterlogging conditions. We are further interested to use this information to better predict whole-tree responses and inform management of trees in urban environments.
We are interested to address this knowledge gap and better understand how roots mediate plant responses to drought and waterlogging conditions. We are further interested to use this information to better predict whole-tree responses and inform management of trees in urban environments.
Building the Fine-Root Ecology Database

Trait-based approaches provide a framework to investigate plant strategies for resource acquisition, growth, and competition, as well as plant impacts on ecosystem processes. Despite significant progress capturing trait variation within and among stems and leaves, identification of trait syndromes within fine-root systems and between fine roots and other plant organs is limited. This empirical roadblock is also partly responsible for the highly simplistic and descriptions of fine-root and belowground ecology in terrestrial biosphere models. To address the need for a comprehensive root trait database, we have compiled the Fine-Root Ecology Database (FRED) from published literature and unpublished data.
With continuing support from the Department of Energy’s Office of Biological and Environmental Research, our collaboration with Oak Ridge National Laboratory and Colleen Iversen has led to the creation of the largest global database of root traits. The roughly 170 root traits currently housed in FRED are described in detail here. For more information visit the FRED Website. These efforts are ongoing as we work to incorporate additional data streams, both new and old, and facilitate even greater ease of use for this freely available, public resource. Despite its young age, FRED has already led to new analyses and insights into fine-root ecology and now serves as the backbone of a new, internationally funded working group linking root traits to ecosystem function (sROOT funded by iDiv).
Figure (Iversen and McCormack unpublished). Map of locations associated with trait observations in the Fine-Root Ecology Database (FRED).
Relevant Publications
Caplan JS, SJ Meiners, H Flores-Moreno, ML McCormack. Fine-root traits are linked to species dynamics in a successional plant community. Ecology, 100: e02588 (2019).
McCormack ML, AS Powell, CM Iversen. Letter: Better plant data at the root of ecosystem models. Eos, (2018).
McCormack ML, D Guo, CM Iversen, W Chen, DM Eissenstat, CW Fernandez, L Li, C Ma, Z Ma, H Poorter, PB Reich, M Zadworny, AE Zanne. Building a better foundation....New Phytologist, 215: 27-37 (2017).
Iversen CM, ML McCormack et al.... A global Fine-Root Ecology Database to address belowground challenges in plant ecology. New Phytologist, 215: 15-26 (2017).
Freschet GT, OJ Valverde-Barrantes, CM Tucker, JM Craine, ML McCormack et al... Climate, soil and plant functional types as drivers of global fine-root trait variation. Journal of Ecology, 105: 1182-1196 (2017).
With continuing support from the Department of Energy’s Office of Biological and Environmental Research, our collaboration with Oak Ridge National Laboratory and Colleen Iversen has led to the creation of the largest global database of root traits. The roughly 170 root traits currently housed in FRED are described in detail here. For more information visit the FRED Website. These efforts are ongoing as we work to incorporate additional data streams, both new and old, and facilitate even greater ease of use for this freely available, public resource. Despite its young age, FRED has already led to new analyses and insights into fine-root ecology and now serves as the backbone of a new, internationally funded working group linking root traits to ecosystem function (sROOT funded by iDiv).
Figure (Iversen and McCormack unpublished). Map of locations associated with trait observations in the Fine-Root Ecology Database (FRED).
Relevant Publications
Caplan JS, SJ Meiners, H Flores-Moreno, ML McCormack. Fine-root traits are linked to species dynamics in a successional plant community. Ecology, 100: e02588 (2019).
McCormack ML, AS Powell, CM Iversen. Letter: Better plant data at the root of ecosystem models. Eos, (2018).
McCormack ML, D Guo, CM Iversen, W Chen, DM Eissenstat, CW Fernandez, L Li, C Ma, Z Ma, H Poorter, PB Reich, M Zadworny, AE Zanne. Building a better foundation....New Phytologist, 215: 27-37 (2017).
Iversen CM, ML McCormack et al.... A global Fine-Root Ecology Database to address belowground challenges in plant ecology. New Phytologist, 215: 15-26 (2017).
Freschet GT, OJ Valverde-Barrantes, CM Tucker, JM Craine, ML McCormack et al... Climate, soil and plant functional types as drivers of global fine-root trait variation. Journal of Ecology, 105: 1182-1196 (2017).