Module I · The Apprenticeship

How a Plant Works

Before you can keep a plant alive, you need to know what it's actually doing.
How a Plant Works

This is the foundation of the whole Apprenticeship. Ten short lessons, about an hour of reading plus a 45-minute field exercise, take you from “I water it when the leaves droop” to a working mental model of the plant as a system. Every later module is operating instructions for the system you learn here.

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Sources & further reading

Brundrett, M. C., & Tedersoo, L. (2018). Evolutionary history of mycorrhizal symbioses and global host plant diversity. New Phytologist, 220(4), 1108–1115. https://doi.org/10.1111/nph.14976

Carter, R., Woolfenden, H., Baillie, A., Amsbury, S., Carroll, S., Healicon, E., Sovatzoglou, S., Braybrook, S., Gray, J. E., Hobbs, J., Morris, R. J., & Fleming, A. J. (2017). Stomatal opening involves polar, not radial, stiffening of guard cells. Current Biology, 27(19), 2974–2983.e2. https://doi.org/10.1016/j.cub.2017.08.006

De Schepper, V., De Swaef, T., Bauweraerts, I., & Steppe, K. (2013). Phloem transport: A review of mechanisms and controls. Journal of Experimental Botany, 64(16), 4839–4850. https://doi.org/10.1093/jxb/ert302

Dunn, B., Singh, H., & Leckie, R. (2017). Mycorrhizal fungi (HLA-6449). Oklahoma State University Extension. https://extension.okstate.edu/fact-sheets/mycorrhizal-fungi

Evert, R. F. (2006). Esau’s plant anatomy: Meristems, cells, and tissues of the plant body (3rd ed.). Wiley. https://doi.org/10.1002/0470047380

Feng, Q., Zhu, S., Wang, X., Liu, Y., Zhao, J., Dagdas, Y., & Nowack, M. K. (2026). Root hair lifespan is antagonistically controlled by autophagy and programmed cell death. Nature Plants, 12, 1022–1033. https://doi.org/10.1038/s41477-026-02279-8

Fenster, C. B., Armbruster, W. S., Wilson, P., Dudash, M. R., & Thomson, J. D. (2004). Pollination syndromes and floral specialization. Annual Review of Ecology, Evolution, and Systematics, 35, 375–403. https://doi.org/10.1146/annurev.ecolsys.34.011802.132347

Hetherington, A. M., & Woodward, F. I. (2003). The role of stomata in sensing and driving environmental change. Nature, 424(6951), 901–908. https://doi.org/10.1038/nature01843

Klatt, B. K., Holzschuh, A., Westphal, C., Clough, Y., Smit, I., Pawelzik, E., & Tscharntke, T. (2014). Bee pollination improves crop quality, shelf life and commercial value. Proceedings of the Royal Society B, 281(1775), Article 20132440. https://doi.org/10.1098/rspb.2013.2440

Koch, G. W., Sillett, S. C., Jennings, G. M., & Davis, S. D. (2004). The limits to tree height. Nature, 428(6985), 851–854. https://doi.org/10.1038/nature02417

Lawson, T., & Blatt, M. R. (2014). Stomatal size, speed, and responsiveness impact on photosynthesis and water use efficiency. Plant Physiology, 164(4), 1556–1570. https://doi.org/10.1104/pp.114.237107

McCree, K. J. (1971). The action spectrum, absorptance and quantum yield of photosynthesis in crop plants. Agricultural Meteorology, 9, 191–216. https://doi.org/10.1016/0002-1571(71)90022-7

Michaels, T., Clark, M., Hoover, E., Irish, L., Smith, A., & Tepe, E. (2022). The science of plants: Understanding plants and how they grow. University of Minnesota Libraries Publishing. https://open.lib.umn.edu/horticulture/

Mississippi State University Extension Service. (n.d.). Can I use a vibrating toothbrush rather than the expensive electric pollinators? Retrieved August 11, 2026, from https://extension.msstate.edu/agriculture/crops/commercial-horticulture/greenhouse-tomatoes/can-i-use-vibrating-toothbrush-rather-the-expensive-electric-pollinators

Morard, P., & Silvestre, J. (1996). Plant injury due to oxygen deficiency in the root environment of soilless culture: A review. Plant and Soil, 184(2), 243–254. https://doi.org/10.1007/BF00010453

Nemali, K. (2018). Understanding the pores of a soilless substrate (HO-287-W). Purdue University Extension. https://www.extension.purdue.edu/extmedia/HO/HO-287-W.pdf

Ollerton, J., Alarcón, R., Waser, N. M., Price, M. V., Watts, S., Cranmer, L., Hingston, A., Peter, C. I., & Rotenberry, J. (2009). A global test of the pollination syndrome hypothesis. Annals of Botany, 103(9), 1471–1480. https://doi.org/10.1093/aob/mcp031

Paauw, M., van Hulten, M., Chatterjee, S., Berg, J. A., Taks, N. W., Giesbers, M., Richard, M. M. S., & van den Burg, H. A. (2023). Hydathode immunity protects the Arabidopsis leaf vasculature against colonization by bacterial pathogens. Current Biology, 33(4), 697–710.e6. https://doi.org/10.1016/j.cub.2023.01.013

Pedersen, O., Sauter, M., Colmer, T. D., & Nakazono, M. (2021). Regulation of root adaptive anatomical and morphological traits during low soil oxygen. New Phytologist, 229(1), 42–49. https://doi.org/10.1111/nph.16375

Pundt, L. (2017). Pollination of greenhouse tomatoes. University of Connecticut Extension. https://ipm.cahnr.uconn.edu/wp-content/uploads/sites/3216/2022/12/2017pollinationofgreenhousetomatoes-1.pdf

Purcell, L. (2021, August 24). How do trees use water? Purdue Landscape Report, 21-14. https://purduelandscapereport.org/article/how-do-trees-use-water/

Raven, P. H., Evert, R. F., & Eichhorn, S. E. (2013). Biology of plants (8th ed.). W. H. Freeman.

Roldán Serrano, A., & Guerra-Sanz, J. M. (2006). Quality fruit improvement in sweet pepper culture by bumblebee pollination. Scientia Horticulturae, 110(2), 160–166. https://doi.org/10.1016/j.scienta.2006.06.024

Singh, S. (2016). Guttation: Mechanism, momentum and modulation. The Botanical Review, 82(2), 149–182. https://doi.org/10.1007/s12229-016-9165-y

Smith, S. E., & Smith, F. A. (2011). Roles of arbuscular mycorrhizas in plant nutrition and growth: New paradigms from cellular to ecosystem scales. Annual Review of Plant Biology, 62, 227–250. https://doi.org/10.1146/annurev-arplant-042110-103846

Taiz, L., Zeiger, E., Møller, I. M., & Murphy, A. (2015). Plant physiology and development (6th ed.). Sinauer Associates.

Terashima, I., Fujita, T., Inoue, T., Chow, W. S., & Oguchi, R. (2009). Green light drives leaf photosynthesis more efficiently than red light in strong white light. Plant and Cell Physiology, 50(4), 684–697. https://doi.org/10.1093/pcp/pcp034

Tyree, M. T., & Zimmermann, M. H. (2002). Xylem structure and the ascent of sap (2nd ed.). Springer. https://doi.org/10.1007/978-3-662-04931-0

Urban, J., Ingwers, M. W., McGuire, M. A., & Teskey, R. O. (2017). Increase in leaf temperature opens stomata and decouples net photosynthesis from stomatal conductance in Pinus taeda and Populus deltoides × nigra. Journal of Experimental Botany, 68(7), 1757–1767. https://doi.org/10.1093/jxb/erx052

Venturas, M. D., Sperry, J. S., & Hacke, U. G. (2017). Plant xylem hydraulics: What we understand, current research, and future challenges. Journal of Integrative Plant Biology, 59(6), 356–389. https://doi.org/10.1111/jipb.12534

Zhang, Z., Zhu, L., Li, D., Wang, N., Sun, H., Zhang, Y., Zhang, K., Li, A., Bai, Z., Li, C., & Liu, L. (2021). In situ root phenotypes of cotton seedlings under phosphorus stress revealed through RhizoPot. Frontiers in Plant Science, 12, Article 716691. https://doi.org/10.3389/fpls.2021.716691