·arno ~a Volume G 1· Number 4· 2002 Page 2 Gestalt Dendrology: Looking at the Arnoldia (ISSN 004-2633; USPS 866-100) is Whole Tree published quarterly by the Arnold Arboretum of Peter Del Tredici Harvard University. Second-class postage paid at Boston, Massachusetts. 9 John Adams, Farmer and Gardener Corhss Knapp Engle Subscriptions are $20.00 per calendar year domestic, $25.00 foreign, payable m advance. Single copies of 15 The Discovery and Rediscovery of the most issues are $5.00; the exceptions are 58/4-59/1 Horse Chestnut (Metasequoza After Fifty Years) and 54/4 (A Source- H. Walter Lack book of Culuvar Names), which are $10.00. Remit- tances may be made m U.S. dollars, by check drawn 20 The Handsome (and Useful) Horse on a U.S. bank; by international money order; or Chestnut by Visa or Mastercard. Send orders, remittances, Klaus K. Loenhart change-of-address notices, and all other subscription- related commumcations to: Circulation Manager, 23 The Horse Chestnut: Accolades from Arnoldia, The Arnold Arboretum, 125 Arborway, Charles S. Sargent Jamaica Plain, Massachusetts 02130-3500. Telephone 617/524-1718; facsimile 617/524-1418; 25 Index to Volume 61 e-mail [email protected]. Front cover A species of Cecropia growing m Postmaster: Send address changes to Equador’s Amazon Basm, clearly showmg the tree’s Arnoldia Circulation Manager modular construction. As trees age, the modules that The Arnold Arboretum define their architecture repeat themselves, becoming 125 Arborway smaller and more numerous. All cover photographs Jamaica Plain, MA 02130-3500 by Peter Del Tredici Inside front cover The trunk of a bnstlecone pme, Karen Editor Madsen, Pmus anstata, growing on Mt. Evans m the Rocky Mary Jane Kaplan, Copyeditor Mountams accomodatmg the presence of a dead branch. Andy Wmther, Designer Inside back cover Environmental stresses can have a Editonal Committee profound influence on the form of a tree. In the case Phylhs Andersen of this Gmkgo biloba, those stresses anse not only Ellen S. Bennett from nature but from humans. Robert E. Cook Back cover An unusual case of nutnent recycling m Peter Del Tredici red oak, Quercus rubra Originally, a wound was Gary Koller covered with callus growth that eventually pushed its Stephen A. Spongberg way inside the rotting wood of the trunk. The callus Kim E Tnpp tissue formed a mushroom-like structure to fill the cavity, which eventually developed adventitious Copyright © 2002. The President and Fellows of roots. These roots are clearly "recycling" nutnents Harvard College from its own moist, rooting heartwood. Gestalt Dendrology: Looking at the Whole Tree Peter Del Tredici he most remarkable thing about trees is they develop and change over time. howEvery year they add height and girth in a flush of new growth. They are forever expand- ing, from the bottom to the top and from the inside to the outside-a tree that is not expand- ing is a tree that is dying. The growth of trees is totally different from that of vertebrate animals, which tend to reach their full developmental potential relatively early in life, and then main- tam themselves in the mature stage for as long as possible. To put it another way, animals are closed and entire in their development while trees are open and expansive. The easiest way to visualize what is meant by open development versus closed is to look at the different approaches to dealing with bodily injury. In trees, if a limb is broken off, then so be it; the trunk will grow around the break and attempt to cover over the dead branch by pro- ducing callus tissue that grows inward from the outer edges of the wound. In many cases, the tree will also produce a new branch just below the location of the old one. Some people call this response wound-healing, but in reality the tree is simply walling off the damaged or dead tis- sue-compartmentalizing it-in an effort to protect the undamaged portions of the trunk. Wound healing revealed in the trunk of an Dead tissue embedded within the trunk is of umdentified comfer that is supportmg a porch m downtown Alaska. The naked wood little consequence so long as rot does not spread Skagway, shows the that into the living wood (Shigo 1986). Mammals, of clearly complex mterlockmg gram forms where the branch is attached to the trunk, cannot tolerate the of dead tis- course, presence as well as where wound-healmg callus tissue has sue within their bodies. If they are to survive, been produced after the branch died. they must repair the injured body part; growing a new one is not an option. three scientists have defined the field, tree archi- Understanding what trees are and how they tecture describes the processes that regulate grow is central to the discipline known as tree the growth and development of trees. Contrary architecture, which was developed durmg the to the meaning of the term architecture when 1970s by a Frenchman, Francis Halle, a Dutch- applied to buildings, tree architecture is about man, Roulof Oldeman, and an Englishman (and dynamic change in tree form over time; it is not Harvard professor), Barry Tomlinson. As these about the static, geometrical shapes of mature The distmctme, "layered" architecture of the pagoda dogwood, Cornus controversa, growmg at the Arnold Arboretum. This type of model is displayed by many understory trees found m temperate, deciduous forests. 4 trees seen in field guides. To put it another way, tree archi- tecture describes how trees develop their shapes, not what shapes they display. The discipline is particularly exciting because it deals with trees holistically, as intact, well-integrated organisms that are greater than the sum of their parts. Compared to most modern biology with its fixation on DNA sequencing, tree architecture takes a refreshingly nonreductionist approach to development. The Meristem Meristem is the term used for the specialized tissue that allows trees to continue expanding throughout their entire life span. This mer- istematic tissue produces the leaves, the stems, the flow- ers, the bark, and the roots- the differentiated tissues of the plant-while remaining undifferentiated itself. What makes meristematic tissue unique is that it exists in a perpetually embryonic state that allows the tree to be reborn every spring through- out its entire life. Four different types of mer- istems are produced by trees. The most obvious one is the Tree is with meristematic centers shoot meristem, or growing growth contmuously embryonic, locahzed m the shoot and root and m the vascular and cork point, which is located at the tips cambmms. Clockmse from top left: heart of bud on every every Expandmg buds of Fraser’s magnoha, Magnolia frasem, showmg tree. On a mature specimen promment, fohaceous stipules that protect the expandmg leaves. All leaf oak, for example, there can be tissue, as well as floral, is produced by the shoot menstem. thousands, if not tens of The pmmary and lateral roots of the red mangrove, Rhizophora thousands of buds, each with mangle. The brackish water the tree grows in makes it easy to observe the the root a tiny dome of branchmg of system embryonic The cambium the tissue that is the meristem. layer of japanese maple, Acer palmatum, made msible by the growth of wound-mduced callus a botched Shoot meristems followmg produce attempt at graftmg. and leaves, flowers, primary The dramatic, exfohatmg bark of the paperbark maple, Acer gmseum. twigs. Located at the tip of The more extensive the cambium growth, the more extensive the exfohation. 5 every root-and again, there are thousands of root tips on a mature tree-is a root meristem that, over time, produces the tree’s massme, underground root system. The third type of meristem is the vascular cambium, a column of tissue that sheaths the trunk, the branches, and the roots of the tree, and is responsible for the secondary increase in girth m all these parts. The vascular cam- bium is basically a gigantic cylin- drical meristem that outlines the periphery of the entire tree and produces the wood that forms the bulk of the tree. When as that broken branch mjury-such A fence post trom lodgepole pme, Pmus contorta, m Yellowstone mentioned the earlier-exposes National Park, clearly showmg the vamation m annual growth rmgs. cambium layer to the elements, it is the vascular cambium that produces the cal- prints of vascular cambium activity, with each lus tissue that overgrows the wood. The last ring accurately recording the amount of growth a type of meristem found m trees is the cork cam- tree makes in any given year. Because tree bium, which produces the bark that protects growth is mostly a function of rainfall, the width and insulates the tree. In general, the growth of of an annual growth ring provides an indirect the cork cambium keeps pace with the growth measure of the moisture available to the tree of the vascular cambmm, sloughing off the old that year. Scientists have used the information layers of bark as it produces new ones. embedded m the width of growth rings to recre- Whereas the specialized cells that compose ate past rainfall patterns that go back, quite liter- meristems retain their full developmental ally, thousands of years (Cohen 1998). Similarly, capacity throughout the life of the tree, cells in for trees that shed portions of their bark in dis- mammals can express their full developmental crete plates, such as london planes, stewartias, potential only in very young embryos. As the and ponderosa pines, the size of the plates that mammalian embryo ages, a liver cell can only are sloughed off each year serve as an mdirect produce a liver cell and a brain cell a brain cell.