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2.1 The Puzzle of Inheritance   17


                       worth no more than the cost of wool, meat, and skin.”   Figure 2.5  Mendel’s garden and microscope. (a) Gregor
                       Which would it be? According to the meeting’s recorded   Mendel’s garden was part of his monastery’s property in Brünn.
                       minutes, current breeding practices offered no definite an-  (b) Mendel used this microscope to examine plant reproductive
                       swers. In his concluding remarks at this sheep-breeders   organs and to pursue his interests in natural history.
                       meeting, the Abbot Cyril Napp pointed to a possible way   (a): © Biophoto Associates/Science Source; (b): © James King-Holmes/Science
                                                                           Source
                       out. He proposed that breeders could improve their ability
                       to predict what traits would appear in the offspring by find-
                       ing the answers to three basic questions: What is inherited?
                       How is it inherited? What is the role of chance in heredity?
                          This quandary is where matters stood in 1843 when
                       21-year-old Gregor Mendel entered the monastery in
                       Brünn, presided over by the same Abbot Napp. Although
                       Mendel was a monk trained in theology, he was not a rank
                       amateur in science. The province of Moravia, in which
                       Brünn was located, was a center of learning and scientific
                       activity. Mendel was able to acquire a copy of Darwin’s On
                       the Origin of Species shortly after it was translated into
                       German in 1863. Abbot Napp, recognizing Mendel’s intel-
                       lectual abilities, sent him to the University of Vienna—all
                       expenses paid—where he prescribed his own course of
                       study. Mendel’s choices were an unusual mix: physics,
                       mathematics, chemistry, botany, paleontology, and plant
                       physiology. Christian Doppler, discoverer of the Doppler
                       effect, was one of his teachers. The cross-pollination of
                       ideas from several disciplines would play a significant role
                       in  Mendel’s  discoveries.  One  year  after  he  returned  to
                       Brünn, he began his series of seminal genetic experiments.
                       Figure 2.5 shows where Mendel worked and the micro-
                       scope he used.

                                                                           (a)
                       Mendel Devised a New Experimental
                       Approach

                       Before Mendel, many misconceptions clouded people’s
                       thinking about heredity. Two of the prevailing errors were
                       particularly misleading. The first was that one parent con-
                       tributes most to an offspring’s inherited features; Nicolaas
                       Hartsoeker, one of the earliest microscopists, contended in
                       1694 that it was the male, by way of a fully formed homun-
                       culus inside the sperm (Fig. 2.6). Another deceptive notion
                       was the concept of blended inheritance, the idea that pa-
                       rental traits become mixed and forever changed in the off-
                       spring, as when blue and yellow pigments merge to green
                       on a painter’s palette. The theory of blending may have
                       grown out of a natural tendency for parents to see a combi-
                       nation of their own traits in their offspring. While blending
                       could account for children who look like a combination of
                       their parents, it could not explain obvious differences be-
                       tween biological brothers and sisters nor the persistence of
                       variation within extended families.
                          The experiments Mendel devised would lay these
                       myths to rest by providing precise, verifiable answers to the
                       three questions Abbot Napp had raised almost 15 years ear-
                       lier: What is inherited? How is it inherited? What is the   (b)
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