Teaching Mendelism

W I L L I A M D . STANSFIELD

H-l

regor Mendel (1822-1884) is rightly credited as being the "father of modern genetics." He presented the results of his pea experiments at a meeting of his local natural history society in two lectures during 1865. His paper was published in the proceedings of the society the next year. From his breeding experiments wilh the edible pea, he recognized the phenomena of what we today call dominance and recessiveness, segregation of aleles, independent assortment of different traits, equal parental contributions lo offspring, and several others. In this article, 1 present some information that might be helpful in two primary respects for those who teach genetics: (1) Teacher Preparation, and (2) Teaching Techniques,

Teacher Preparation
The topics discussed in this segment are intended to broaden the knowledge of teachers regarding parts of Mendel's 1866 paper that have been subject lo dispute and seldom appear in biology textbooks. They may help prevent teachers from making statements that Mendel did not make or possibly that he did not mean to infer. Also, Mendel made several major contributions to the study of heredity oiher than his famous "rules" or "laws" that are not commonly mentioned in most biology textbooks. Whether or not these topics should he presented to a class depends in part on the educational (cognition) level of the students, but they are entirely at the discretion of the teacher. Most geneticists have little time or inclination to delve into the liisiiiry of their discipline because they are concerned with the latest research- In fact, very jew people have read Mendel's paper and, tjmorig those who have, very few have understood it. Coreos and Monaghan, 1993 Few teachers have had time and/or easy access to the original publications (or translations thereof) that form the basisof each of the disciplines we are assigned lo teach. What many of us know about Mendelism has been obiaincd from secondhand sources, which may contain interpretive errors and/or omissions of important facts. Though popularly cited as being one of the most excellent research papers in nineteenth-ceniury biology, it is not one that 1 would recommend to be on a reading lisl for advanced biology students or biology teachers without first being made aware of the followmg facts. One problem is that Mendel wrote in German. The title of his paper was Versuche ber Pflanzen-Hybriden (see reference MendelWcb for the original German paper) or "Hxpcriments on Plant Hybrids." Most Americans are not riiieni enough in German lo translate his paper for ourselves. WiuiAM D. STANSREW, Ph.D., is Emeritus Professor, Biological Sciences Department, California Polytechnic Stale Universiiy, San Luis Obispo, CA 93407; e-mail: wstansji&calpoly.edu.

Translations sometimes fail to express the thoughts or intents of the author accurately from one language to another. Just two English iranslalions of Mendel's 1866 paper may have been the main sources of most textbook information. The first translation was commissioned by the Royal Horticultural Society (RHS) in 1901 (Mendel, lB66a). The second translation (Mendel, 1866b) was made by Eva Sherwood who stated that this more recent translation was made because a "careful comparison with the original German text showed not only ,i number of mistakes which fundamentally changed the mean ing of Mendel's sentences but in addition so many other inai curacies ..-". This is the version ihai 1 have used in preparinii most of the present article. For those who wish to researcii Sherwood's translation of Mendel's paper, 1 have added pag. numbers in parentheses for each of Mendel's quotes in tin following text. Mendel used some terms that may be subject to interpretation. "The only fault, which occurs on several occasions in the Versuche, is that the term iraif (Merkmal) is used to mean either phenotypc or aii-f, depending on ihe context, whL-h indicates that the distinction may not always have been clear or entirely sharp in Mendel's own mind" (Hartl & Orel. 1992). Furthermore, he did not use many of the terms thai are now so familiar to us, including gene, genetics, aleles, gametes, zygote, somatic cell, chromosome, nucleus, haploid, diploid, homozygous, heterozygous, genotype, phenotype, F|, and F2. It is awkward (and probably largely inappropriate at the pre-college level) for teachers to try to explain Mendel's work using his own words, so most teachers use these modern terms even though they do not appear in his famous 1866 paper.

U H

What Did Mendel Set Out To Discover?

Many textbook authors claim that Mendel set out [ discover the basic rules of heredity. But according to Mendel (p.12), his purpose was "to determine the number of different forms in which hybrid progeny appear, permit cla.^ sification of these forms in each generation with certainty, and ascertain their numerical interrelationships." Some ABT readers may disagree with the following view, but according to Coreos and Monaghan (1993), Mendel's data have been widely interpreted as being about heredity, but they were not interpreted in that way by him. The word "inheritance" appears only once in Mendel's 1866 paper (p, 12) wbere he discusses the variable intensity of the green color of cotyledons and concludes that this phenomenon is "not inherited by the offspring. ' The science of Mendelian genetics devei oped after 1900 as the 1866 paper was reinterpreted by oili ers in light of what had been learned about reproduction anil cytology since its publication.
V-

What Was Known Before Mendel?

Before presenting the essence of Mendel's contributions (Mendelism), it should first be emphasized to students that

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in MendeVs day many biologists believed that hereditary material was a fluid, and trans acquired by an individual during its lifetime could be transmitted to progeny (Lamarckism). Nothing was known about genes, chromosomes, mitosis, or meiosis, Mendel was aware that other biologists had reported examples of plant hybrids "that remain constant in iheir progeny and propagate like pure strains ... This feature is of particular importance to the evolutionary history of plants hecause constant hybrids attain the status of new species" (p. 41 ). Mendel made no mention of Darwin's theory of evolution hy natural selection, but he "studied the German translation of Charles Darwin's On the Origin oj Species, published in 1863" (Orel. 1984). For lack of any other theory, Darwin provisionally accepted Lamarckian Inheritance as a possible mechanism for producing heritable variations on which natural selection could operate when he wrote his On the Origin oj Species (first published in 1859). There is no evidence that Darwin ever read Mendel's paper. The fact that Mendel was searching for a mechanism whereby new species couid originate might suggest that he probably was an evolutionist and not just a plant breeder. But at least one source presents a contrary view. "Darwin's concepts were continuous variation, mutation, and 'soft' heredity [acquired characters; Lamarckism]; Mendel espoused discontinuous variation and 'hard' [materialistic, atomistic] heredity without mutation" (Bishop, 1996). The orthodox doctrine of special creation, to which the Catholic monk probably subscribed, denied the existence of constant hybrids. Mendel also knew that pea flowers have female and male reproductive organs enclosed in a structure called the "keel," formed by the union of three of the five petals, which insures natural seli-fertilization. He knew how to remove the anthers from a flower hefore its own pollen is shed, and how to transfer pollen from another plant onto the stigma to create hybrid seeds. He knew the role of ovules (Mendel called them "germinal cells") and pollen cells in fertilization, "IP]ropagation in phanerogams [an outmoded term referring to the Spermatophyta, characterized by the production of pollen tubes and seeds; a]] angiosperms and gymnosperms] is initiated hy the union of one germinal cell and one pollen cell..." (Mendel 1866h, p, 41). And he knew that there often was no sharp distinction between species and varieties of a species (p. 5).

one alele to each endosperm cell. A single dominant alele (A) governing an endosperm trait (e.g., yellow color) wii] cause the endosperm to deve]op the dominant trait even in the presence of two recessive aleles {cia) for green color. Thus, if somatic cells of the seed parent are genetically aa. and those of the pollen parent are AA. the endosperm of the resulting seed will be yellow and genetically aaA. Mendel states that the shape of seeds and the color of seed albumen "develop immediately after artificial fertilization merely through the influence of the foreign pollen. Therefore they can be observed in the first year of experimentation, while the remaining traits do not appear in the plants raised from fertilized seeds until the following year" (p. 10). I believe, for the reasons stated above, that Mendel cou]d not have known how many copies of a gene were represented in various plant parts and/or at various times in the plant's life cycle. Not everyone agrees with me. Here is the prohletn, Mendel often uses a single letter (A) to represent the breeding structure of a plant that produces, upon selfing, only plants with the dominant trait; the lower case letter (a) represents a plant that breeds true for the alternative recessive trait; and the two letters (Aa) represent a plant that segregates progeny with both dominant and recessive traits. On one occasion, Mendel (p. 30) presents the fonnula:
A/A * A/a + a/A + a/a = A + 2A + a.

What Was Not Ktiown in Mendel's Time?

All biology teachers know ihai double fertilization occurs in flowering plants. One sperm nucleus unites with (fertilizes) the egg nucleus to form the embryo; another sperm nucleus unites with two polar nuclei within the embryo sac (ovule) to form the endosperm. What they ail may not know is thai this was discovered in 1898 by S. G. Navashin, 14 years after Mendel's death. Mendel bad a microscope, but it is doubtful that he had seen the growth of a pollen tube down the style or the union of nuclei during fertilization. Even if he had seen nuclei, he might not have thought that his hereditary elements were confined therein rather than being distributed throughout the cell. Mendel probahly knew that cotyledons are the leaf-forming parts of the embryo in a seed. They function as storage organs from which the seedling draws food, or they may absorb and pass on to the seedling nutrients stored in the endosperm (called "albumen" by Mendel). Pea seeds thus have no endosperm at maturity. Once the cotyledons are exposed to light, they develop chlorophyll and function as the first eaves of a plant. Mendel did not know that the egg nucleus is haploid (n). ceils of the embryo are dipioid (2n), and endosperm cells are triploid (.3n). The maternal (seed) parent or the ovule contributes two identical aleles to endosperm. The paternal (pollen) parent contributes only

In discussing hybrids of species other than Pisurn, Mendel (p. 41) states that "'|I]t seems permissible to assume that the germ cells of tbose [hybrids] that remain constant are identical, and also like the primordial cell [zygote?] of the hybrid," For HartI and Orel, tbis clearly implies that the homozygous forms A and a each contain two hereditary determinants, "The key question is whether the word identical (gleichartig) is intended to mean 'identical in number' or 'identical in type,' We presume tbat Mendel meant identical in both senses" (Hartl & Orel, 1992). However, it appears to me that, for the sake of simplicity (Occam's razor; Stansfieid, 2002), Mendel assumed that for each trait a minimum of two genes was present in its somatic ceil genotype, and only one gene was present in gametes. He did not prove that only one gene of a gene pair is in a gamete or that onv two aleles are present in the genotype of somatic cells. It might have occurred lo his audience that the results of his experiments would not have changed if gametes carried two or more identical copies of a gene. As long as dominant and recessive aleles segregate in the production of gametes, the numher of gene copies in a gamete or in a genotjq^e of an embryo or somatic eel] is not critical. Recall that endosperm contains three haploid sets of chromosomes (triploid, 3n), two sets of maternal origin and one set of paternal origin. One dominant and two recessive aleles [Aaa) produce the same endosperm phenot>'pe as two dominant and one recessive alele i^AAu). The same principle would apply to somatic cells regardless of their ploidy state. Lesl we he too harsh in our anachronous criticism oj Mendel's somewhat incoiisistent use ojsymhoh, it is worthwhile to bear in mind that modern Drosophila gent-iiclsts routinely use unpaivcd symbols when rejcrring to homozygous rcessives; jor example, in rcjerring to Drosophila strains, the symbol al means the genotype al/al, and cn bw meiitis the genotype cn bw/cn bw. Hartl and Orel, 1992 Mendel may have used the term "antagonistic elements" (p. 42) to represent what are now known as "aleles." Some readers of Mendel's paper have had difficulty interpreting him where he states (p. 43): "In the formation of [reproductive] cells

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all elements present participate in completely free and uniform lashion, and only those |elemenis] that differ separate from each other" (tialits added). Some say ihai ihis violates one of the lencts of Mendclian genetics, viz. "the law of segregation." Today we know ihal identical aleles in homozygotes also segregate during meiosis and the formation of gametes. Mendel provided for the segregation of the characters of the hybrid in gamete formation, but did not refer this to segregation of determiningfactors. Since there are no characters to segregate in gamete formation in the true-breeding lines, he did not provide for segregation of anything in these cases. His noi doing so is further evidence that he was ifjin^ with empirically determined things, and not with theoretically postulated determiners. Monaghan and Coreos, 1985 The integuments are maternal tissue surrounding the ovule. They become the seed coat. "The angiosperm seed is consequently a 'genetic mosaic' consisting of maternal, zygotic, and endospermal tissue, each having its own chromosomal and genetic constitution" (Swanson, 1957). Given these facts, we would expect that the color of the seed coat would be determined by the genotype of the motber plant, not by the genotype of the seed embryo or endosperm. It is not clear tbat Mendel understood this fact. Nevertheless, Mendel reported (p. 10) that the color of the seed coat is correlated with tbe color of the flower in which it developed. Plants bearing violet-red flowers produce seed coats witb grey-brown coats; whiie-ilowered plants produce white seed coats. The grey-brown or leather-brown color of tbe seed coat is also associated with reddish spots on tbe leaf axils. Textbooks tbat ciic Mendel as tbe father of tbe "one-gene, one-trait hypothesis" ignore tbe fact that Mendel was aware that some genes are associated with muliiple pbenoiypic effects. It is not as widely known as it sbould be that Mendel's 1866 paper contains two major segments. Tbe first part deals with peas and alternative qualitative "characters of kind" exhibiting disconiinuous variation, typical of classical Mendclian traits such as red flowers vs. white flowers. Tbe second part of Mendel's paper deals with hybrids of other plant spectes and "characters of degree" exhibiting continuous variation typical of morphometric (quantitative) traits such as sizes and shapes of various body parts. Mendelian traits involve only one or a few genes witb major pbenoiypic effects (oligogenes) and are relatively easy to study and explain. The phenotypic variation of quantitative traits, on the other band, is commonly governed by many genes (polygcnes), eacb with relatively small but cumulative effects on tbe phenotj-pe tbat may vary from one set of environmental conditions to anoiber. Tbese facts considerably complicate tbe study of this class of traits. Mendel discussed the polygenic trait of llower color in beans. He even offered a two-gene model to explain the spectrum of colors that can segregate from bean hybrids. This aspect of Mendel's contributions is neglected in sotne textbooks. It is interesting to note thai Mendel studied two discontinuously varying traits in peas that usually are under polygenic control in other plant species - length of stem (tall vs. short) and shape of seed (round vs. wrinkled). In the first part of his paper, Mendel uses tbe term "factors" (p. 24) which has been widely interpreted by others as tbe equivalent of what we would today call "genes." He first uses tbe term "elements" (p. 42) in the second segment of his paper instead of the word "factor." It is not known why he changed tbese terms in these two segments of his paper, but it only serves to confuse the reader. "Nowhere in the Versuche is the physical nature [particles, fluids, emulsions] of die Elemente [the elements] discussed in

enough detail to infer bow Mendel might have imagined them. ... It does not matter whether Mendel was thinl<ing in terms of particles or lluids, since he emphasized repeatedly ibe key point that differing elements emerge unchanged from their association |in hybrids]" (Hartl & Orel, 1992).

Why Was Mendel's 1866 Paper Ignored for so Long?

Mendel's paper had no perceivable influence on the study of heredity until 1900 when it was independently discovered by tbree others. Why was ibis so? Some people tbink tbat his work was ignored because Mendel was a monk with no scientific credentials. Some thought that bis paper was not distributed widely enougb throughout Europe. Others suggest that scientists were too occupied with the ramifications of Darwin's evolutionary theories and the prevailing paradigm of the inheritance of acquired characteristics. Still otbers think that Mendel's use of probability theory was foreign to solving biological problems or too cotnplex for biologists to understand- "[Mendel's] mathematical analysis of trails probably would not bave made sense lo anybody but mathematicians - who probably would not have had the least bit of interest in pea plants" (Starr fi Taggart, 1981 ). 1 think that the main reason why Mendel's work was not appreciated in bis time was because it was premature. Had bis audience known about cbromosomes. mitosis, and meiosis, they migbt have been better prepared to accept Mendel's tben radical ideas. Without this knowledge, his audience might bave had many questions that Mendel was probably unable to answer, such as those in the following list. What arc the chemical and physical characteristics of Mendel's hereditary factors? By what mechanism(s) is segregation of aleles accomplis bed? How is the phenomenon of dominance explained physiologically and/or developmen tally? When, in the plant's life cycle, do genes replicate and by what mecbanism is this accotnplished? How many copies (replicas) of a gene are in gametes or somatic cells? Does segregaiion of aleles occur in bomozygous plants as well as in hybrids? Do all somatic cells contain the same gene composition and number of gene copies, or are some genes depleted as cells differentiate to form various parts of the plant? What implications, if any, does Mendel's work have for Darwin's theory of evolution by natural selection? With so many questions left unanswered, it is little wonder that Mendel's work was so unappreciated during bis lifetime. Moreover, Mendel could find no e\idence that hybrid peas, when self-fertilized, breed true to type, i.e., hybrids thai produce only hybrid progeny without segregating any pure bomozygous types- Hence, bis pea hybrids would not be considered new species by Lhe criteria of his day. Those who were hoping thai Mendel's experiments might shed light on tbe origin of species via hybridization must have been disappointed.

Teaching Techniques
According to modern educational philosophy, some of our most important missions as science teachers are to give our students the opportunity to solve problems, analyze empirical

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dala, formlale testable hypoiheses to explain the data, and think critically. Part of Mendel's genius was lhe way he designed and carried oui his experiments. Students should at least be asked to think about experimental designs as a critical factor to the success or failure of any scientific research. For example, "How would you design an experiment lo find out if red or white flower colors are inherited trails or if these traits are induced by environmental conditions? Why did Mendel choose peas for his experiments rather than some other plants? Because time is sueh a limiting factor in our classrooms, we may feel compelled to initially discuss cell structures and functions, mitosis, and meiosis prior to presenting the results of Mendel's experiments and his interpretations thereof. But there is no "right way" or "wrong way" to teach genetics. For example, 1 found one biology textbook (Marslan, 1964) that presents the genetics of feather color in Andalusian fowl before presenting any of Mendel's pea experiments. The three feather colors (black, blue, and white) are explained as being governed by a pair of codominant aleles, so that allelic segregation in heterozygous parents is expected to produce in their progeny three phenot>'pes in the ratio 1:2:1, respectively. This may be an easier way for some students to grasp the essentials of genetics belore discussing Mendel's results with its complicating phenomena of dominant and recessive aleles (Allchin, 2000). I would like to suggest a different approach that attempts to satisfy as many of the aforementioned lofty educaiiona! objectives as possible. Students could initially be told what biological knowledge Mendel had (or could have knowTi) and what he could not have known at the time he was doing his breeding experiments. They should also he made aware of what theories of heredity were popular in his time. Then the students could be asked to try to place themselves in the audience when Mendel presented the results (raw data) of his experiments, but prior to his analysis and interpretation of the data. Each student would then try to explain these results, given the state of knowledge at the time. This exercise helps students appreciate lhe historical perspective of how advances in scientific knowledge have occurred. It also requires the applications of critical thinking and problem solving. If students do not suggest reducing the raw data to their lowest whole number ratios, the teacher should ask them to do so. For example, Mendel reported that hybrid parents produced 705 progeny with red flowers and 224 with white [lowers; thus, 705/224 = 3:L5:1 ratio. Remember, Mendel was well schooled in chemistry and physics and was used to analyzing experimental data in this way (Monaghan & Coreos, 1983). Once this is done, students should be asked to interpret ihese ratios. If they find it difficult to respond, the teacher could prompt them by asking questions such as: "What do the data suggest about the nature of the hereditary substances (fluids vs, particles) and/or the theory of acquired characteristics? If hereditary substances were fluids like red and white paints, what color would mixtures (hybrids) of two diffrent paints be expected to produce?" Tbereafter, Mendel's interpretation of the data can be divulged by the teacher and discussed by the classStudents then could be asked to concentrate on any subjects that Mendel might nol have mentioned or discussed adequately. In other words, at the end of Mendel's lectures, what questions might his audience have liked to ask him? A starter list of such questions has been provided above. After a list of these kinds of questions have been raised by the class, students should be allowed to let their imaginations soar in their attempts to provide plausible (potentially tesiable) answers (hypothesis construction), even using their own terms if they wish (as long 348 THE AMERICAN BIOLOGY TEACHER, VOLUME 70, NO. 6, AUGUST 2008

as they are adequately defined). For example, when attempting to explain how segregation of aleles might occur mechanistically, perhaps someone may suggest that allelic particles of ihe same type might have an affinity for one another and thus tentl to clump together, as in the formation of a crystal. Alternatively, if dominant and recessive ailelic particles are of dilferent sizes, a molecular sieve of some sort might be able to separate them. Can different liquids spontaneously segregate into homogeneous regions? Have you ever tried to mix oil and waier? Il doesn't matter that these ideas are not the "true" explanation for segregation of aleles. The main ihing is lo get the students involved in some "thinking outside the box" (beyond wbat is presently known). This may be a more important aspect of advancing science than anything else in the scientific method. Now, if it has not been done before, this would be an appropriate time to explain DNA structure, chromosomes, mitosis and meiosis. With this new knowledge, the results of Mendel's experiments and his principles of allelic segregation and independent assortment ot different traits become more easily understood. By having teachers, and students aitkally analyze the state oj science at the time of Mende this may perhaps offer greater insight into how dramatic and revolulionary MendeVs findings were. In addition, the interpretations of Mendel's findings by students in context of the science of the time might spark interest in the students for further study of the work of Mendel. Then as students c^nilnuf on their foray into modern genetics, they might have a better understanding of how crucil and signi/ifn ihe body of Mendel's scientific work was. Anonymous reviewer of a previous draft of this paper Ideally, we might like to use this instructional technique as widely as possible. Realistically, however, considering the breadth of subjects biology teachers are required to present, we can employ this technique in relatively few cases. But ii we could use il in only one subject area of biology, where else would ii apply better than when we are teaching Mendelism?

Epilogue
During neither my undergraduate nor graduate education was I required to read Mendel's 1866 paper. Sometime after 1966, I did read Sherwood's translation of it, but not criiically I didn'i completely understand parts of Mendel's paper, bui 1 thought it was largely due to my misunderstanding of some of his terms. It wasn'l until sometime after 1993, when 1 read the book by Coreos and Monaghan (1993), that 1 began lo learn biology textbooks might not be accurately presenting what Mendel was thinking, I had retired from teaching by iben, so I was not able to enlighten my students with this new perspective. However, after retirement, I soon began work on a book (Stansfield, 2000) for a general audience in which I discussed some of the problems biologists have had interpreting Mendel's work. Over the years since then, 1 have continued to research and ruminate on how I might use this knowledge to best assist biology teachers. In this article, I present some of the results of this process. 1 therefore would appreciate hearing from educators who have tried to leach Mendelism along the lines suggested here, so that I might assess and report (to ABT) whether or not it has been belpful in meeting the educational objectives listed previously.

References
Allchin, D.K, (2000)- Mending Mendelism. The American Biology Teacha\ 62(9),633-639.

Bishop, B.E. (1996). Mendel's opposition to evolution and to Darwin. Journal aJ Heredity. 87. 205-213. Coreos, A.F, & Monaghan, EW. (1993), Gregor Mendel's Experiments on Plant Hybnds - A Guided Study. New Brunswick, NJ: Rutgers University Press. Darwin. C ( 1859), On the Origin of Species by Means of Natural Selecon of Favoured Races in the Strugglefor Life. London:John Murray. Hartl, D.L. &r Orel, V. (1992). What did Mendel think he discovered? Genetics, JJi, 245-253. Marslan, D. ( 1964). Principles of Modem Biology, 4th Edition. New York: Holt, Rinehari and Winston, Mendel, G. ( 1866a). Experiments in plant hybridization. English translation made by the Royal Horticultural Society of London with footnotes added and minor changes suggested by Professor W. Bateson, enclosed within [ |. In J.A. Peters (Ed.) (1959). Classic Papers in Genetics (pp. 1-20), Englewood Cliffs, NJ: Prentice-Hall. A complete copy of this English translation is available at the Web siie: http:// www, niende] web, org/ M ende.l, hjnil. Mendel, G. (1866b). Experiments on plant hybrids. English translation made by E, Sherwood, In C, Stern and E, Sherwood (Eds.)

( 1966). The Origin o/Genetics; A Mendel Source Booh (pp. 1-55). San Francisco: W.H. Freeman and Company. McndelWeb, This Web site contains a copy of Mendel's original paper in German. Available online al: http://www.mendelweb, org/MWGerText.html. Monaghan, F.V. & Coreos, A.F. (1983). Possible influences of some 19th century chemical concepts on Mendel's ideas about heredity. Journal oj Heredity. 7t 297-299. Monaghan, F.V. & Coreos, A.F. (1985), Mendel, the empiricist, Joumdl of Heredity. 76. 49-54. Orel, V. (1984). Mendel. New York: Oxford University Press. Stansfield, W.D. (2000). Death of a Rat: Understandings and Appreciations of Science. Amherst, NY: Prometheus Books, Stansfield, W.D. (2002), Occam's razor and the nature of scientific theories. The Amccan Biology Teacher. 64(2), 107-109, Starr, C. &Taggart, R. (1981). Bioh^: The Unity and Di\ersity of Life, 5lh Edition. Belmont. CA: Wadsworth Publishing Company, Swanson, C.P (1957). Cytology and Cytogenetics. Englewood Cliffs, NJ: Prentice-Hall,

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