Hereditas He compared the investigation of Mendel’s research to the In his book, “The origin of genetics: a Mendel source book,” Stern. THE LIFE OF GREGOR JOHANN MENDEL ‐TRAGIC OR NOT? ÅKE GUSTAFSSON. DEPARTMENT OF FOREST GENETICS, STOCKHOLM AND INSTITUTE. View Hereditas dalam hukum from ENGLISH at Sampoerna University. 1. Tanaman ercis dipilih dalam percobaan Mendel karena memiliki.

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In this article, alternative explanations for this close agreement that inheritance in pea does not conform to kendel standard statistical model, that data were omitted, that ambiguous data were categorized to better match predicted ratios, and that some data were deliberately falsified are tested using approaches that are designed to distinguish between these alternatives.

The possibility that garden pea Pisum sativum L. Considerable evidence is introduced that Mendel omitted some of his experimental results, but this alternative cannot adequately explain the low average deviation from expectations that is characteristic of the segregation data he presented.

It has been years since Gregor Mendel published a brief article presenting compelling evidence for the then novel concept that individual heritable units control specific traits. Some science historians have mrndel that he really does not deserve this title as his article emphasized the behavior of plant hybrids and he never fully described the 2-allele genetic system that we are familiar with today Corcos and Heredihas ; Monaghan and Corcos Fisher argued that if Mendel planted only 10 seeds per family the actual expected ratio would be 1.

Others have suggested that it is more likely that Mdndel planted more seed from each F 2 plant but included only 10 in the data he reported Wright ; Orel and Hartl ; Fairbanks and Rytting Normally, in a scientific article, a strict interpretation of the text would be appropriate, and Fisher insisted on this interpretation. Most of the data that Mendel presented in his article are summarized in Table 1. Furthermore, the data are not subdivided into units that maximize the degrees of freedom in the results but rather are presented by trait in order to facilitate comparisons with data generated on that same trait by other researchers.

Because of the debate mentioned above regarding the number of seeds actually planted by Mendel for his testing the 2: Taking each trait separately, the segregation ratios for the 3 traits with only one entry each pod color, pod shape, and stem fasciation do not appear to be questionable on their own.

However, I suspect Fisher would still question the close agreement between observed and predicted ratios, particularly those for the seed characters.

Beyond the possibility that Mendel fabricated at least some of his data, there are several alternative hypotheses, listed below, that have been suggested by Sturtevant as well as by others. In the following 4 sections I will examine each of the alternatives, using data from a considerable number of studies in pea published between and to test the plausibility of each of the alternatives.

As unlikely as this possibility may seem, it has been given some attention in the past and it also provides an opportunity bereditas introduce aspects of pea biology that are directly relevant to segregation data in this species. It was postulated by Thoday that pollen tetrad formation in pea could proceed in such a way that in a heterozygous plant pollen grains possessing alternate alleles were somehow grouped so that the process of pollination was not a random sampling of pollen grains but directed to enforce a consistent 1: Although it is doubtful that Thoday or others seriously supported this hypothesis, it has been tested in several ways as the correlated pollen model Seidenfeld or tetrad-pollen model Fairbanks and Schaalje and rejected as a possibility.

Another method of testing the general possibility that pea gives more consistent segregation ratios than predicted by statistical models is to examine the segregation data in pea published by other researchers.

The results of the 7 post-Mendel determinations were similar to those of Mendel in that none gave a significant deviation from the expected 3: F 2 segregation data from several early 20th century geneticists testing pea cotyledon color for conformance to a 3: One limitation to the studies listed by Johanssen is that all were done to directly test the hypothesis that traits give a 3: Thus, all the investigators hededitas be accused of having a bias similar to that herfditas of Mendel: To address this limitation, I have assembled additional data reported in the literature by a number of pea researchers who published later than those Johannsen cited Table 3 and Supplementary Table S1.

The difference between the data sets listed by Johannsen and those in Table 3 and Supplementary Table S1 is that the latter contain only data that were part menndel linkage studies. In such studies, the investigator is not as interested in testing segregation ratios as in classifying the segregating traits accurately so that linkage between traits can be identified.

The data were selected to include only those traits also investigated by Mendel and involve population sizes comparable with those Mendel used population sizes less than were not included.

The reason for assembling the linkage data into 2 tables is that many crosses within what is currently recognized as Pisum sativum L. Mendel screened his crosses for high fertility. However, for completeness these data sets are presented in Supplementary Table S1. Table 3 presents 30 linkage segregation data sets for A testa and flower color24 for Le stem height33 for Gp pod color28 for R seed shape15 for I cotyledon color6 for Fa flower distribution on stemand 6 for V pod shape.


Including data sets from Supplementary Table S1 only increases the differences between the linkage data and those of Mendel. To summarize, rather than indicating that pea segregation data is likely to conform more closely to expected ratios, the result from the linkage studies suggests the opposite.

In such a graph, the expected frequency is equal for all units along the x axis. The distribution of the studies cited by Johannsen appears to fit this prediction, although the sample size is too small to provide a significant test.

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hereditxs Neither the data of Mendel nor those of the linkage studies provide a good fit to the predicted distribution. Thus, even after elimination of results that might be attributed to wider crosses than Mendel used, the data from linkage studies do not support the alternative currently being discussed—segregation ratios produced in pea do not tend to give higher P values than would be predicted by standard statistical models. The units on the vertical axis are the proportion of data sets in the respective P value range for the specified source of data.

Fisher discounted this alternative because of the limited garden space Mendel had available to grow his plants, making it unlikely that Mendel could have grown many more plants than he mentioned in his manuscript.

Hence, based on space availability and statistical arguments, it does not appear that Mendel could have grown a sufficient number of populations to select only those that would provide the particularly low variance and close agreement to his model. It is doubtful that in his studies Mendel encountered as much segregation distortion as is present in Table 3 and Supplementary Table S1, but he should have observed some.

He selected 22 lines for his crossing studies, and described them as being from 4 different species P. Even within this subspecies there exist lines that when crossed will produce F 2 progeny with distorted segregation ratios Kosterin and Bogdanova This possibility is further supported by the fact that in his analysis of hybrids segregating for several factors, Mendel observed that 2. Thus, particularly in his bi- and trifactorial crosses, Mendel probably crossed lines with relatively divergent genetic backgrounds, comparable to many of the crosses used to produce the data in Table 3.

The conclusion relevant to the current discussion is that although Mendel screened his hybrids and discarded those showing a significant loss of fertility, he still encountered some F 2 populations that gave weak plants. He probably eliminated some F 2 populations that displayed a considerable fraction of weak plants, but he did not have time or space to eliminate a significant proportion of his populations. Thus, even in his relatively limited number of analyses, Mendel should have observed some distorted sets.

The data in Table 3 reveal that for over half the mapping experiments cited the segregation at this locus was distorted. In the crosses giving distorted segregation at this locus in Table 3all give an excess of the wild-type phenotype, consistent with incomplete penetrance of the mutant phenotype. Mendel examined over F 2 individuals for this trait, which probably reflects the progeny from at least 30 hybrid plants.

Undoubtedly, the line s he used as the mutant parent was highly penetrant for the trait. However, once Mendel crossed the trait into a mixture of genetic backgrounds, it is doubtful that the trait would always exhibit high penetrance. For fasciated stem it seems most likely that Mendel obtained his data from only 1 or 2 crosses that gave excellent penetrance of the trait. If so, there should have been data from other crosses involving Fa that displayed skewed ratios that were not included in his article; otherwise he could not have concluded that this trait assorted independently from all others.

As Mendel acknowledges directly in his manuscript that wide variation could occur in observed segregation ratios, there appears no reason for him to eliminate the relatively few cases that failed to support his hypothesis. His method of presenting the bulk of his data combining data from several crosses into 1 segregation set, such as for pod color, pod shape, and fasciation would allow him to selectively exclude certain data sets from a segregating population while including others.

Just how thoroughly data need to be presented in a manuscript is somewhat subjective, as is evident to anyone who has gone through preparation and review of a scientific manuscript.

Gregor Mendel’s classic paper and the nature of science in genetics courses.

Such selection of data could have eliminated divergent segregation sets and allowed Mendel to slightly reduce the overall variance in his data.

However, herecitas would not be sufficient to bring into doubt his general conclusions, and it hegeditas doubtful that just elimination of data could explain the absence of segregation results with P values less than 0.

As will be discussed later, it is likely that Mendel simply eliminated highly divergent results because he did not want to confuse his audience with experimental details that detracted from menvel demonstration of his model.

This alternative was suggested by both Fisher and Sturtevantbut a test of this hypothesis has yet to be performed. To undertake such a test we need to know more about the phenotypes Mendel was trying to classify in order hereitas establish which traits might produce such uncertain or intermediate phenotypes.


The exceptional property of pea that made it so valuable to Mendel was its suite of polymorphic morphological characters that turned out to be primarily controlled by single loci in the germplasm he used. Thus, near the very start of his manuscript Mendel dismisses the possibility that errors in classification of phenotypes occurred.

He does not mention further problems with phenotypes and when he presents his experimental data he does not indicate encountering ambiguous phenotypes or discarding data that he thought might be suspect. Notably, in his description of the dihybrid and trihybrid crosses where he details the number of seeds that did not germinate and the number of plants that failed to reproduce, there is no mention of plants expressing uncertain phenotypes.

The reader must accept that all plants were completely and accurately classified. However, in my own experience, and I believe for many other individuals who have analyzed these traits, the phenotypes of several of the traits are not always clearly discernible despite considerable familiarity with their variation.

Tall plants also have longer internodes and often have longer peduncles and tendrils.

Gregor Mendel’s classic paper and the nature of science in genetics courses.

Indeed, scoring segregation of this gene is usually performed by classifying flower color rather than seed coat color. The yellow pod character produces an upper stem, peduncles, and pedicels with a yellowish tinge. In addition, the yellow pods typically have thinner pod walls.

Mendel mentioned many of these associated traits and presumably could have used them to confirm instances where there may have been some doubt about the primary character. A much more detailed description of the variation in seed shape can be found in Khvostova An excellent method of discriminating between round and wrinkled peas is to examine the shape the starch granules in the cells of the cotyledons. This feature was used by several of the researchers cited by Johannsen but was unknown at the time Mendel did his studies.

Pod shape and flower distribution on the stem are 2 additional traits that tend to be difficult to score in certain crosses, although these were not mentioned specifically by Mendel as problematic. The edible podded character results from a reduction of schlerenchyma in the pod wall. This reduction can be produced by either of 2 unlinked mutations, p or vthe pattern of the reduction of the schlerenchyma being different in the 2 mutations Rasmusson Due to its slightly greater amount of schlerenchyma, the pod wall in the latter mutation does not always collapse between seeds and is often harder to distinguish from the wild type Rasmusson Yet even Rasmusson, who worked extensively with vdid not appear to be able to consistently score the phenotype in various genetic backgrounds, determining linkage frequencies between V and Le ranging from 5 to 15 cM Rasmusson The 2 loci are close together at one end of pea linkage group III, and the variance in the linkage intensities determined by Rasmusson probably reflects more the difficulty in classifying pod phenotype than a variation in recombination frequency.

As has been mentioned above, the phenotype and penetrance of the fasciated stem mutation varies in different genetic backgrounds. We are faced with the possibility that either he selected 1 cross that gave him good fertility and an excellent segregation ratio at Fa or much of the data is fabricated.

I feel there is sufficient concern about the Fa data that I have omitted them from the following analyses. If there were a tendency to classify dubious individuals in a way to hereitas the expected ratio, segregation ratios for these traits should fit the expected ratio more closely than the data obtained for traits with unambiguous phenotypes.

Two approaches were used to evaluate this possibility. Theoretically, the number of data sets in each bin pair should be equal. Chi-square analysis based on the predicted 1: As the data sets are correlated the same plants are used for the 3 ambiguous traits, the Bonferroni correction should be used, reducing the critical P value 3-fold e.

Comparison of P value distributions derived from the segregation of Mendel hereeditas those of the linkage studies cited in Table 3. The Mann—Whitney U test Mann and Whitney was also employed to compare the range of P values in ambiguous and nonambiguous traits. For the data from the linkage studies there was no significant difference in P value distributions between the 2 categories.

Thus, both approaches indicate that a confounding influence was acting when seed shape, pod shape, and cotyledon color were being classified. The most straightforward explanation is that the investigator, whether it was Mendel or an associate, being faced with having to classify ambiguous phenotypes did so with a mnedel to what was predicted by the model.

The possibility of an assistant deliberately altering the data to appease Mendel has been mentioned in the literature herediats times.

However, Fisher believed that Mendel had developed his model before starting any experiments, and thus his expectations would have been known to assistants before much data were collected. It is plausible that Mendel assigned the classification of seed phenotypes hereditsa others while handling the vegetative characters stem height, pod color, pod shape, and flower distribution menndel.