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6 Black Walnut CHAPTER 6 6 Black Walnut Charles H. Michler, Keith E. Woeste, and Paula M. Pijut USDA Forest Service, Hardwood Tree Improvement and Regeneration Center at Purdue University, 715 West State Street, West Lafayette, IN 47907-2061, USA e-mail:[email protected] 6.1 hydroxy-1, 4-naphthoquinone), naturally occurring Introduction in the leaves, buds, bark, nut husks, and roots of black walnut, is a highly selective, cell-permeable, ir- 6.1 .I reversible inhibitor of the parvulin family of peptidyl- Origin and History prolyl cis/trans isomerases (PPIases) and functions by covalently modifying sullfhydryl groups in the tar- Black walnut (Juglans nigra L.), also known as eastern get enzymes (Henning et al. 1998; Chao et al. 2001). black walnut or American walnut, is a fine hardwood Certain plants, especially tomato, apple, and several species in the family Juglandaceae, section Rhyso- conifer species, are adversely affected (allelopathy; caryon (Manning 1978). In general, J. nigra will not foliar yellowing, wilting, and even death) by being cross with species in the sections Cardiocaryon or grown near the roots of black walnut trees (Good- Trachycaryon, but J. nigra will cross with J. ailan- ell 1984; Dana and Lerner 1994). Horses can con- tifolia (Cardiocaryon) (Williams 1990). Juglans nigra tract acute laminitis, an inflammation of the foot, will also hybridize to some extent with other Juglans when black walnut wood chips or sawdust is used species (Dioscaryon and Rhysocaryon), and one hy- for stall bedding or stables and paddocks are located brid is recognized: J. nigra x J, regia = J. x interme- too close to walnut trees (Galey et al. 1991). Histor- dia Carr. (USDA-NRCS 2004). Native to the decidu- ically, the bark of black walnut was used by several ous forests of the eastern United States (USA), from Native Americans, including the Cherokee, Delaware, Massachusetts to Florida and west to Minnesota and Iroquois, and Meskwaki, in tea as a cathartic, emetic, Texas, and occurring naturally in southern Ontario, or disease remedy agent, and chewed or applied for Canada, black walnut is seldom found in pure stands, toothaches, snake bites, and headaches (Moerman but rather in association with five mixed mesophytic 1998, 2003). Caution: the bark should be used cau- forest cover types: sugar maple, yellow poplar, yel- tiously in medicine because it ispoisonous. The Chero- low poplar - white oak - northern red oak, beech kee, Chippewa, and Meskwaki also used the bark to - sugar maple, and silver maple - American elm make a dark brown or black dye (Moerman 1998, (Williams 1990). Black walnut is a large tree and on 2003). The Comanche pulverized the leaves of black good sites may attain a height of 30 to 38 m and di- walnut for treatment of ringworm, the Cherokee used ameter of 76 to 120 cm and can exceed 100 years of leaves to make a green dye, and the Delaware used age (Williams 1990; Dirr 1998; USDA-NRCS 2004). the leaves as an insecticide to dispel fleas (Moerman Black walnut is shade intolerant, and control of com- 1998, 2003). The nut meats were also a food source peting vegetation is especially important in new plan- for Native Americans, and the nuts are still consumed tations for the first 3 to 4 years. Black walnut grows today by people and are an important food source for best on moist, deep, fertile, well-drained, loamy soils, wildlife. although it also grows quite well in silty clay loam soils or in good agricultural soils without a fragipan (Williams 1990; Cogliastro et al. 1997). These sites 6.1.2 include coves, bottomlands, abandoned agricultural Botany fields, and rich woodlands. Black walnut forms a deep taproot, wide-spreading lateral roots, and has been Juglans nigra (section Rhysocaryon, Fig. 1) is the cultivated since 1686. A toxic chemical 'juglone' (5- largest and the most valuable timber tree of theluglans Genome Mapping and Molecular Breeding in Plants, Volume 7 Forest Trees C. Kole (Ed.) O Springer-Verlag Berlin Heidelberg 2007 190 C. H. Michler, K. E. Woeste, P.M. Pijut Fig. 1. Form, leaf shape, bud (~haracters, and flower and fruit morphology of Juglans ni- gra L. " Copyright 2001 - Bruce Lyndon cu&nGgham species and is hardy to USDA hardiness zone range or in clusters of two to three and are edible, sweet, of four to nine (Dirr 1998). Black walnut is monoe- oily, and high in protein (Reid 1990). The nut is sub- cious with male and female flowers maturing at dif- globose to globose, rarely ellipsoid, 3 to 4cm, and ferent times (McDaniel 1956). Staminate catkins (5 to very deeply longitudinally grooved, and the surface 10 cm) develop from axillary buds on the previous between the grooves is coarsely warty (Flora of North year's wood and appear as small, scaly, conelike buds, America Editorial Committee 1993+). Leaves are al- and the female flowers occur in two- to eight-flowered ternate, pinnately compound, 30 to 60 cm long, with spikes borne on the current year's shoots (Brinkman 9 to 23 leaflets, nearly glabrous and somewhat lus- 1974; Williams 1990; Flora of North America Edito- trous dark green, pubescent and glandular beneath, rial Committee 1993-t; Dirr 1998). The female flowers with petioles 6.5 to 14 cm long covered with glandu- more commonly appear first (protogyny) and flow- lar hairs (Flora of North America Editorial Commit- ering occurs with or shortly after the leaves. Because tee 1993+; Dirr 1998). Black walnut stems are stout, flowering is dichogamous, self-pollination is unlikely, densely grey-downy, smooth, and reddish buff, and which promotes outcrossing. The fruit is a drupe- they have a chambered light brown pith (paler than like, furrowed nut enclosed in a thick, indehiscent that of butternut, J. cinerea) and a distinctly notched yellowish-green husk that develops from a floral in- leaf scar. Terminal buds are ovoid or subglobose, 8 volucre (Brinkman 1974). Fruits are subglobose to to 10 mm long, and weakly flattened (Flora of North globose, rarely ellipsoid, 3.5 to 8 cm, warty, with scales America Editorial Committee 1993+). Lateral buds and capitate-glandular hairs (Flora of North America are smaller, often superposed, and greyish in color. Editorial Committee 1993+). The fruit occur singly J. nigra has a dark grey or brownish bark, deeply split Chapter 6 Black Walnut 191 into narrow furrows and thin ridges, and the ridges (USDA-FAS 2004). Black walnut yields edible nuts are chocolate in color when cut, forming a roughly that are used in baking (cookies, cakes, etc.) and ice diamond-shaped pattern. The sapwood of black wal- cream products. The Hammons Products Company nut is nearly white, and the heartwood varies from (Stockton, MO; http://www.black-walnuts.com)is the light to dark brown. The wood is heavy, hard, strong, world's premier processor and supplier of American stiff, normally straight grained, and has good resis- black walnuts for both food and industrial uses. Se- tance to shock (Forests Products Laboratory 1999). lection of black walnut trees for nut quality and pro- The chromosome number of black walnut is 2n = 32 duction has developed slowly over the years, but over (Woodworth 1930). Black walnut trees produce seed 700 cultivars have been named and the percent of edi- at about 12 years of age, with good seed crops oc- ble kernel has improved to over 34% (Reid 1990; Reid curring every 2 to 3 years (Brinkman 1974). Seeds of et al. 2004). Black walnuts are low in saturated fats black walnut, like most Juglans spp., have a dormant (3.4 g per 100 g edible nut), have zero cholesterol, and embryo, but dormancy can be broken by fall sowing are high in polyunsaturated (35.1 g per 100 g edible or by moist prechilling of seeds at 1 to 5 "C for 3 to nut) and monounsaturated fats (15 g per 100 g edible 4 months (Brinkman 1974). nut) (USDA-ARS 2004). Black walnuts are also a good source of protein (24.1 g per 100 g edible nut) and fiber (6.8 g per 100 g edible nut) containing low levels of 6.1.3 sugar (1.1 g per 100 g edible nut) (USDA-ARS 2004). Economic Importance Ground black walnut shell (see Hammons Products Company, http://www.black-walnuts.com)is a hard, Black walnut is one of the largest hardwood trees durable, nontoxic, biodegradable abrasive product found in the USA and is valued economically and used for blast cleaning and polishing. It is also used for ecologically for its wood and edible nuts. Owners industrial tumbling and deburring, as well as for uses of quality black walnut wood receive high market in oil-well drilling, water filtration, and as explosive prices, and the wood has many uses including fur- fillers. niture, veneer, cabinets, interior architectural wood- work, flooring, and gunstocks. Black walnut wood with figured grain gets even higher market prices. 6.2 Curly and wavy figure can produce interesting char- acteristics in veneers, and these can arise from walnut Black Walnut Genetics butts, crotches, and burls. The nut is an important food source for wildlife and is also consumed by hu- Black walnut genetic research has been focused on mans. The majority of black walnut trees occur in the practical improvement of the species for the pro- natural stands, with walnut plantations (ca. 13,800 ac) duction of timber or nuts. Although black walnut has accounting for 1% of all the black walnut volume (ft3) a large native and commercial range, black walnut in the USA (Shifley 2004). There are 11 states that improvement has been largely a Midwestern preoccu- currently have the greatest volume of black walnut pation.
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