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Excitability in Plant Cells Author(S): Randy Wayne Source: American Scientist, Vol Sigma Xi, The Scientific Research Society Excitability in Plant Cells Author(s): Randy Wayne Source: American Scientist, Vol. 81, No. 2 (March-April 1993), pp. 140-151 Published by: Sigma Xi, The Scientific Research Society Stable URL: http://www.jstor.org/stable/29774870 Accessed: 06-04-2016 10:15 UTC REFERENCES Linked references are available on JSTOR for this article: http://www.jstor.org/stable/29774870?seq=1&cid=pdf-reference#references_tab_contents You may need to log in to JSTOR to access the linked references. Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at http://about.jstor.org/terms JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. Sigma Xi, The Scientific Research Society is collaborating with JSTOR to digitize, preserve and extend access to American Scientist This content downloaded from 130.223.51.163 on Wed, 06 Apr 2016 10:15:49 UTC All use subject to http://about.jstor.org/terms Excitability in Plant Cells An external stimulus to a plant, such as touch, can trigger a cellular mechanism that generates a defensive response Randy Wayne cells, in fact, are hotbeds of electrical ac? ber of ions crossing the membrane. As a duck a pond, paddles it nips alongat the topsthe ofedge under? of tivity, and plant studies have provided They concluded that ions carry the cur? water vegetation. When one nip catches much of the foundation of what is rents that create the action potential. a shoot of Cham, a relative of the green known generally about electrical activi? Although plants are no longer the algae, it sends a spectacular system into ty in cells. Cham has been important in leading organisms used in research on action. The force of the duck's bite trig? those studies and continues to be. the basis of electrical excitability, a num? gers an electrical mechanism in the Physiological studies of electrical ac? ber of investigators have significantly plant, and ionic current rushes across tivity began in the 19th century, and advanced our knowledge of both the the membrane of the nibbled cell. Then since then animal and plant physiolo? mechanisms and the effects of electricity the fluid inside the cell, the protoplasm, gists have worked in parallel. In order in plants. Modern techniques common stops its normal flow around the pe? to study the activity where it happens, to neurophysiology have been applied riphery. The protoplasm quickly jells, at the cellular level, investigators had to to a variety of plants, and the results preventing any leakage that could arise find organisms in which the activity show that electrical physiology in plants from the duck's attack. could be studied in isolation from the is as complex as the systems found in Cham is hardly the only plant that re? whole plant or animal. They also need? animals. Moreover, a variety of plants sponds to external stimuli. All plants re? ed to find cells large enough that they use electricity to initiate action; exam? spond to gravity as they grow, and could be probed with electrodes. In ani? ples are the closing of the leaves of a plants can have various responses to mal studies, the search led to the long Venus flytrap and the touch-driven Hght. Some follow a 24-hour cycle, ad? nerve cells of squids, in which axons, drooping of the leaves of some Mimosa justing the orientation of their leaves for the fibers carrying messages from the species. Nevertheless, the most detailed the maximum absorption of light dur? cell body, are so large that they were information exists for characean algal ing the day. Some plants respond with originally thought to be blood vessels. cells, which I shall examine here. The movement when they are touched by Plant physiologists, on the other hand, electrical activity in these algae is worth predators. selected species of algae that have large examining not only for its importance in What may be less obvious is how cells, such as the characean algae Cham plant biology, but also because studies plants respond to stimuli. Although and Nitella. of plant excitability may help us under? most people know that electrical signals In 1898 Georg H?rmann, a German stand the evolution of the human ner? mediate the responses of an animal's physiologist, observed that big differ? vous system. nervous system, it is less widely known ences in voltage measurements could that plant behavior, too, is governed by develop across cell membranes of Characteristics of Characeans complex electrical mechanisms. Plant Nitella. When such differences are re? Characean algae have been used in generative they are called action poten? much of the work on plant excitability. Randy Wayne received a Ph.D. in botany at the tials, because the regeneration implies They are stoneworts, with a fossil University of Massachusetts at Amherst under the action?the passing of an impulse. By record stretching back to the Devonian guidance of Peter Hepler. Wayne's doctoral work the 1930s, characean algal cells were so period, which began about 400 million considered the contribution of calcium to the phy well known that many investigators years ago, and they are the ancestors of tochrome-mediated signal transduction chain that studied them. For example, K. S. Cole all higher plants. Extant stoneworts be? leads to fern-spore germination. He continued his and Howard Curtis of the National In? long to a single family, Characeae, work on phytochrome as a postdoctoral fellow at the stitutes of Health, who later became which is composed of six genera in? University of Texas with Stan Roux. Wayne began known as pioneers in the electrical ex? cluding Cham and Nitella. The majority his work in membrane biology with Masashi Tazawa citability of squid neurons, began of the extant species inhabit the bottom at the University of Tokyo. He is currently an assis? tant professor of plant biology at Cornell University, studying excitability in Nitella. These in? of clear freshwater ponds, where they where he tries constantly in his teaching and vestigations showed that an action po? live entirely submerged. research to repay the debt he owes to his teachers. tential in Nitella is accompanied by a As I have noted, the primary attrac? Address: Section of Plant Biology, Cornell 200-fold increase in the cell membrane's tion of characean algae as an object of University, Ithaca, NY 14853. conductance, as measured by the num study is the size of their cells. In Chara, 140 American Scientist, Volume 81 This content downloaded from 130.223.51.163 on Wed, 06 Apr 2016 10:15:49 UTC All use subject to http://about.jstor.org/terms Figure 1. Chara, an alga, responds to environmental stimuli, as do many plants. A variety of factors, including mechanical stimulation, can generate an action potential?a transient change in voltage across a cellular membrane?that causes some of this alga's internal fluid to jell, preventing it from leaking through small holes or tears in the plasma membrane. Large cells make Chara an appealing organism for electrical physiology. The plant's shoot is composed of long internodal cells separated by smaller nodal cells, seen here at the tip of a plant (lower right) and supporting reproductive structures (top). Each internodal cell is about six centimeters long and half a millimeter wide; like all plant cells it has three distinct partitions. The outer surface is the cell wall, which is composed of cellulose. Underneath the cell wall is the plasma membrane, which is formed from two layers of lipids. Much of the inside of the cell is taken up by a vacuole, which is bound by the vacuolar membrane. (Photograph at right courtesy of the author.) the plant body is composed of long in? spersed with proteins. Beneath the plas? ternodal cells separated by smaller ma membrane there is a layer of chloro nodal cells. A single internode may be plasts, the sites of photosynthetic six centimeters long and half a millime? processes. Most of the interior of the cell ter wide, or about half as long as a is a vacuole, a sac filled largely with wa? toothpick and half as wide. The inter? ter and bounded by another membrane. nal structure of an internodal cell is un? The area between the vacuolar mem? like that of an animal cell. Like all plant brane and the plasma membrane is cells, the external border is a cell wall, filled with protoplasm; here are found which is composed of cellulose fibers the cell nucleus and the cytoplasm, a that provide rigidity to the cell but are viscous fluid that contains the cell's or permeable to the extracellular fluid. Just ganelles such as mitochondria and ri beneath the cell wall is a semipermeable bosomes. plasma membrane, which is composed The protoplasm of characean cells of two layers of lipids that are inter moves constantly around the periphery 1993 March-April 141 This content downloaded from 130.223.51.163 on Wed, 06 Apr 2016 10:15:49 UTC All use subject to http://about.jstor.org/terms has more positively charged ions and the other side has more negatively charged ions, then there is a potential, or voltage, across the membrane. Here I shall discuss four potentials: membrane potential, resting potential, receptor potential and action potential. A membrane potential is the voltage across a membrane, or a measurement time of the distribution of ions.
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