The Convenience of Arabidopsis Elizabeth A. Kellogg small, it’s plain, it’s absolutely ordinary, but it’s become one of the most popular lab plants around. It’s Arabidopsis thaliana, and its most important characteristic is that it’s handy. It’s Scientists have a long tradition of working with the handy stuff, literally the things close at hand, to answer questions about phenomena that are otherwise inaccessible. Charles Darwin opens The Origin of Species with a chapter that describes in some detail breeding experiments were on pigeons. Pigeons convenient and served as a model. If people could select pigeons for complex characteristics, Darwin reasoned, then could select other organisms the way. Hence, evolution by natural selection: an all-encompassing theory supported in part by experiments on the very nature same plain, thoroughly homely pigeon. Using whatever’s handy is of course an art fully exploited by schoolteachers. If they want to make a mask or a turkey, they use a paper plate. An egg carton does fine for an alligator (or stegosaurus or a bouquet of flowers), and noodles are clearly necklace material. Lab scientists use a lot of those same items. Just like Charles Darwin and schoolteachers everywhere, we work with whatever’s handy. procedure in molecular biology labs, called Southern blotting after the One common who invented it, is carried out in Tupperware containers. Rubbermaid does fine, too; the lid just has to be watertight. man This same procedure can be done with specialized chemicals like dextran sulfate-dextran sulfate being notable because it costs about $400 a pound-but for many purposes ordinary powdered milk from the supermarket works just fine. The standard procedure then involves washing a bit of high-tech nylon membrane in powdered milk in a plastic kitchen container. When the procedure is finished the piece of nylon is wrapped in Saran Wrap. (In fact, in the standard chemical stockroom Saran Wrap is on the shelf right next to all the fancy chemicals.) This procedure is like many others in research; for some purposes only a very particular tool will do (like the special bit of nylon membrane), but there are many cases where you can simply use what’s handy. This same "principle of handiness" applies choosing organisms to study, especially in rapidly growing field of plant molecular biology. People who study plant molecular biology are trying to understand exactly how plants work, down to the details of the DNA to the that make up their genes. Because almost anything they discover will be new, several plants are equally good to begin working on. So why not start with the most convenient? I currently make my living studying genetic relationships in the wheat tribe, a group that Arabidopsis thaliana. Photo and © by Kurt Stepnitz, MSU/DOE Plant Research Laboratory. 14 . includes species, Turkey, barley and rye and a lot of other many of them native to Mongolia, and the Mideast. It happens, how- ever, that three wheat-related species are weeds that grow near the parking lot of the Harvard Bio Labs, and another grows next to the playing fields in my Cambridge neighborhood. It’s obvious which ones I looked at first. As the study has progressed I have had to seek out the less accessible members of the group to fill in the story, but the starting point was arbitrary and determined as much by convenience as by logic. That’s the major attraction of Arabidopsis thaliana to molecular biologists. It does have a common name-mouse ear cress-but it’s rarely used. The plant has no horticultural value. It isn’t edible. But for some purposes it is very convenient, and it has thus become important because of its value as a scientific tool, analogous to the fruit fly [Diosophila). Arabidopsis thaliana is a tiny relative of the cabbage and part of the same family, commonly known as the mustard family and botanically as Cruciferae or Brassicaceae (either name is acceptable). At maturity it is about eight inches tall. It can be germinated by the hundreds on petri plates. Populations of thousands of plants fit easily on one greenhouse bench. The generation time (seed to seed) is about three months. Compare this to that other workhorse of the plant genetics world, maize. Maize plants are large, well over eight inches tall, and planting thousands requires acres of land and scientists with strong backs. The generation time is about six months, but to get two generations a year requires that you plant one winter crop in Hawaii. Arabidopsis clearly has a logistical advantage, although you do lose the excuse to winter in Hawaii. Arabidopsis is also handy in that its internal workings, its genetic machinery, are unusually simple. No extras, no add-ons, no window dressing. One scientist, Dr. Jerry Fink of the Whitehead Institute, has described it as the plant equivalent of a Hyundai. For many studies this makes it the plant of choice. The Boston Area Arabidopsis Group alone includes sixty or so scientists who specialize in topics such as hrp, GA, auxin, rubisco activase, and ribozymes. Most of these formidable-sounding specialities have direct applications to understanding the crop plants that feed humanity. Hrp, for example, is a gene or set of genes involved in the plant’s response to a pathogenic fungus or bacterium. If we know exactly how plants respond to pathogen attack and what allows some plants to resist some pathogens, then we might be able to find ways to reduce our dependence on the toxic chemicals that are now used to control damage by plant pests and to engineer resistant crop plants. GA and auxin are plant hormones that control the rate and timing of growth and development. It’s all basic research, the fertile ground that fosters direct applications. So it is that a modest weed is achieving an eminence formerly reserved for crop plants. There are seedbanks that store Arabidopsis seed, there is an Arabidopsis newsletter and an Arabidopsis Information Service as well as an Arabidopsis Research Initiative. It is becoming, in its own curious way, cally important. economi- There are, of course, questions that A. thaliana can’t help us with. Since we know almost nothing about the relatives of Arabidopsis and very little about its natural history, it is almost useless in studies of evolution within the mustard family. It has, however, become a big part of one story that promises to tell us a lot about the evolution of flowers and their multiple forms. If you look closely at a developing flower during the very early stages when it is best seen with an electron microscope, you will see tiny nubbins for all the floral parts. In Arabidopsis there are four that will be sepals, four that will become petals, six stamen nubbins, and two carpels that will form 15 Scanning electron micrographs of a developing Arabidopsis flower. A. The top of the inflorescence and flower buds. B. A flower bud with one sepal removed to show nubbins that will become stamens; the mound in the center will become the pistil. C. A flower at a slightly later stage. P petal; LS lateral stamen; MS medial stamen ; G gynoecium (pistil). D. A flower bud shortly before openmg. Some of the sepals and petals have been removed. Note that the stamens have not yet elongated fully. Bar 10 mm m A, B, and C; 100 mm thick. Photographs reproduced with permission from Bowman et al. (1 992). = = = = = 16 the two halves of the ovary. Scientists in a California lab headed by Elliot Meyerowitz have found the chemical signals that tell the various nubbins how they should develop. Altering these chemicals can make the stamens turn into petals, or the petals turn into stamens, or even turn all the flower parts into leaves. (This last idea-that flower parts can be viewed as modified leaves-was first suggested in the eighteenth century by the German poet Goethe.) Comparing work on Arabidopsis with studies done on snapdragshows that similar substances appear in other dicot flowers as well. It is now a tantalizing possibility that these chemicals may be involved in generating some of the startling diversity of floral form that we enjoy in our gardens, fields, and forests. Scientific papers are typically written as though the scientist had thought of an unanswered question, carefully designed an experiment, chose a perfect model system, and concluded with a formal and thoroughly rational analysis. (This, incidentally, was not the structure used by Darwin.) Such papers ons fairly easy to read once you get the hang although rarely as pleasurable as The Origin of Species. The style of the scientific paper is simply a late-twentieth-century convention. Unfortunately, it obscures the way that science actually works, the way are of it, decisions are made and directions taken. There is a lot of serendipity involved. There are insights from chance conversations, opportunities created by particular combinations of people, place, and time. And there is the very practical tendency to grasp the tools at hand. Reference Bowman, J. L., H. Sakai, T Jack, D. Weigel, U. Mayer, and E. M. Meyerowitz. 1992. SUPERMAN, a regulator of floral homeotic genes Arabidopsis. Development 114: 599-615 in ol Elizabeth Kellogg is Research Associate in Organismic and Evolutionary Biology at Harvard University and an Associate of the Arnold Arboretum.