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W2430174693 abstract "It's these changes in latitudes, changes in attitudes: Nothing remains quite the same. For all of our running, and all of our cunning: If we couldn't laugh, we would all go insane.—Jimmy Buffett Richards himself expresses surprise that this book has remained popular and in demand more than ten years since publication of the first edition in 1986, which was reprinted in 1990 and 1994. The present revised second edition retains the strengths that drove sales of the original, but it remains a limited and flawed coverage of the dynamic field of plant reproductive biology. That said, I hasten to point out that I have used Richards's treatment of the subject as the basic “textbook” for my own graduate course in Plant Reproductive Ecology and find that it serves the purpose well. Anyone seeking a reasonably comprehensive coverage of this subject would be well served to buy this book. As a “laboratory manual” to supplement Richards's treatment, I use and recommend Kearns and Inouye's (1993) book, Techniques for Pollination Biologists. At first I was struck by the apparent dramatic increase in size of the new edition: compared to the 1990 version, which is only 23.3 mm thick, the 1997 edition is 31.6 mm! Returning to my original 1986 first edition, however, I discovered that it also measured 31.6 mm. Amazingly, all three of these versions contain exactly 529 pages! This is so, despite the fact that Richards deleted more than 300 references and added more than 500 new ones. It does appear to me that the format and production quality have been improved in the second edition, which is printed on “permanent acid-free text paper.” I find it interesting that the term “breeding system” figures so prominently in the title to the book, yet it is not even defined in the glossary! We learn on p. 201 that the breeding system includes, but is not limited to, whether a species is self-compatible or self-incompatible, but the full “suite of characters which control the breeding system, all of which are important, and all of which must be considered together” is never explicated. A list of the major chapter titles is somewhat illuminating in this regard: (2) Sexual theory in seed plants, (3) Sexual reproduction in flowering plants, (4) Floral diversity and pollination, (5) Pollination biology and gene flow, (6) Multi-allelic self-incompatibility, (7) Heteromorphy, (8) Dicliny, (9) Self-fertilization and inbreeding, and (10) Agamospermy. Compared to the first edition, one chapter is deleted entirely: Vegetative reproduction. The other major revisions mentioned by Richards in the preface are expansion of the chapter on sexual theory and that on self-fertilization. Indeed, Chapter 2 (sexual theory) appears to have increased by about 10 pages at the expense of Chapter 3 (sexual reproduction). Chapter 9 (self-fertilization), on the other hand, is within one page of its former length (i.e., not expanded). In fact, the greatest increase in size seems to have occurred in Chapter 10 (agamospermy), which has grown from 33 to 55 pages! I hesitate to criticize an author's coverage of a topic, as I am well aware of what my own biases would be if I were to take on the same daunting challenge. And I feel that, in general, Richards has done an excellent job of covering a wide range of phenomena in a diverse array of plants. His solid grounding in basic genetics, as well as natural history, has stood him in good stead to deal with this material. Nevertheless, there are a number of sins of omission, as well as commission, that I will lay out below for the prospective reader/buyer of this book. I feel that the book is weakened by Richards's decision to delete all of the material on vegetative reproduction. It is a common and widespread phenomenon that is of great importance to plants. If asexual reproduction is to be ignored, then it makes little sense to discuss the issue of “why be sexual?” (p. 24). I also question Richards's claim in the preface that “all references to plant groups other than the flowering plants have also been lost.” On p. 298 he launches a diatribe against applying sexual terminology developed for vascular plants to bryophytes, but his arguments are not developed well. Having considered these issues in some detail myself (Wyatt, 1985), I think Richards should either explain his opposing views more fully or, more propitiously perhaps, omit them here. To me, the greatest weakness of the present volume is Richards's aversion to theory and lack of appreciation of the role of modeling in modern ecological genetics. Richards's lament, voiced throughout the book, is that “theoretical and population studies have grown in popularity” (p. 412) to the detriment of more traditional approaches such as Mendelian genetics, morphology, and natural history. In his view, “theoretical advances are sterile and non-substantive” (p. ix). Moreover, his first edition “was a book based on natural history rather than mathematics, with very little modelling included.” He has attempted to remedy this somewhat in the second edition, but, unfortunately, his efforts have fallen short. He frequently uses ESS (evolutionarily stable strategy) arguments without seeming to understand their basic formulation. His discussion of sexual selection is very confusing and conflates it with gametophytic selection at one point (p. 36). Sometimes, Richards seems to avoid discussing relevant models or statistical approaches (e.g., not explaining mixed-mating models, instead referring the reader to Brown (1990) on p. 370). Other topics are avoided entirely (e.g., “functional gender” is mentioned in passing on p. 25, but nothing of substance is said) or covered very imperfectly (e.g., “neighborhood is a rather difficult concept…” on p. 187). The insights contributed by phylogenetic approaches are not even mentioned! Among the sins of commission committed by Richards in this book, the most glaring are those related to the inclusion of unpublished data and inappropriate rebuttals of the work of others. I have already alluded to the fact that I believe Richards's coverage of agamospermy is overweighed relative to other topics. Much of this derives from the inclusion of unpublished data by Richards and his graduate students, which has never been subjected to peer review. Besides the extensive detailed coverage of unpublished data on Taraxacum (e.g., pp. 349, 399), there are references and new tables and figures relating to flower temperatures in Crocus (p. 108), orchid pollination (pp. 117, 121), fruit predation in Silene (p. 141), and heterostyly and homostyly in Primula (pp. 270, 276, 391). To me, these seem excessive and inappropriate to include in a textbook. Moreover, Richards's critique of Barrett's (1983) review of heterostyly (p. 284) and of an esoteric point regarding agamospermy by two Swedish workers from the 1940s (p. 423) would be more appropriately placed in one of his own papers in the primary literature. Not surprisingly, the author most cited by Richards is Richards: five or more dissertations by his graduate students are cited and at least 21 papers on which he is first author! On a number of subjects Richards's views evince uncritical acceptance of older, mainstream dogmas or, conversely, eccentric rejection of new thinking about a subject. In drawing attention to this, I do not mean to condemn Richards or profess to know that he is “wrong.” Nevertheless, it is important for readers to be aware of these prejudices. On p. 7 Richards baldly states that “most plants have evolved a mixed reproductive strategy.” This view derives from Stebbins (1950) and has been challenged by Lande and Schemske (1985), among others. In Richards's defense, I point out that he does offer a more balanced appraisal of this issue in Chapter 9 (self-fertilization). On p. 12 Richards assumes that gametophytic self-incompatibility is basal within the angiosperms, as proposed originally by Whitehouse (1950). Recent molecular evidence, however, accords more closely with the views of Bateman (1952) and others that self-incompatibility systems have arisen multiple times in various clades. The utility of pollen–ovule ratios as indicators of plant breeding systems seems to be accepted at face value (p. 31), despite serious shortcomings of this approach and questioning even of its conceptual basis by various authors. Richards states that “ explanation that dioecy has arisen from selfing progenitors in response to selection pressures which favour outcrossing has a long and respectable history” (p. 300). This seems to imply that new, ecological explanations, which presumably have a short and disreputable history, should be rejected out of hand. Yet it is exactly this fresh thinking that has breathed new life into a number of well-worn subjects, such as the evolution of dioecy. Finally, Richards's embracing of “agamospecies” (p. 456), which he defines as “microspecies or segregate species of low taxonomic amplitude used in agamic complexes” (p. 456), may grow out of his long-term studies of such plants, but his taxonomic views are distinctly in the minority. There is a serious problem on p. 44 and the following few pages, where the mathematical formulas have been left out entirely. On p. 65 and in the glossary, Richards gives the definitions of “multiovulate” and “uniovulate,” rather than of “multilocular” and “unilocular,” which he intended. On p. 81, it is stated that North American species of Cypripedium produce nectar, but these orchids are actually nonrewarding (Gill, 1989). Richards apparently disagrees with Raven's (1972) view that bird flowers are red because that color makes them less apparent to nectar-stealing bees, arguing instead that birds “are very responsive to red” (p. 96). Mention should at least be made of the alternative hypothesis. In the plate facing p. 244, fig. 1a is not the temperate Passiflora incarnata, but rather the tropical P. vitifolia. To my knowledge and according to several taxonomic treatments that I consulted, Spartina patens is not dioecious, as Richards claims on p. 299. Because I work primarily with mosses and milkweeds, I am, of course, especially sensitive to any mistakes regarding these plants. Apparently, Richards's delimitation of the bryophytes on p. 457 excludes the hornworts (Anthocerotae). I was surprised to learn on p. 130 that asclepiads do not have specialized pollination mechanisms like the orchids. Indeed, in my view, the milkweed floral mechanism actually surpasses the orchids in complexity! Richards concludes that “it is likely that the regular, actinomorphic flowers of the asclepiads preclude the attraction of many more sophisticated visitors such as bees.” This is nonsense, as even a cursory reading of the relevant literature or a short time spent in the field with these plants will attest. This myth is perpetuated later as Richards refers to the “butterfly-pollinated milkweed Asclepias exaltata” (p. 187), which in fact is more often pollinated effectively by bees, as are most milkweeds (including even butterflyweed, A. tuberosa). In the glossary (p. 466), Richards overlooks the fact that milkweeds, as well as orchids, produce pollinia. I enjoyed the rare occasions in this volume when Richards, intended or not, used puns and colorful language to get his point across. For example, on p. 446 it is suggested that sporophytic agamospermy in tropical fruit trees, like Citrus, “is certainly a fruitful field for further investigation.” On p. 87 it was somehow reassuring to learn that pseudocopulatory bees “rarely ejaculate.” I have to admit that it was an issue I would never have thought to explore! A few stylistic and organizational issues are worth considering. I found Richards's frequent interruption of the narrative with numbered lists to be very distracting. I am sure that his intent was to be precise and concise and to facilitate the reader's rapid rediscovery of certain key issues. For me, however, this convention did not work. The lists were overused and sometimes used to summarize points from unpublished manuscripts. The text includes an odd mixture of British and American spelling and word usage. “Hermaphrodity” is used in preference to “hermaphroditism” throughout. On p. 149 and in the glossary on p. 463, the word “major” is explained as a North American word for “graduate.” It actually means to specialize, as in a major field of study, as bees concentrate their foraging on flowers of a single species of plant. I am also unfamiliar with the use of “self-exclusive” (p. 385) to mean mutually exclusive, as hypotheses can be. Nevertheless, the most foreign aspect of Richards's style to me is his predilection for using nouns as verbs or adjectives. Hence, we see “resource” (p. 345) and “compartment” (p. 390) used as verbs, whereas “hermaphrodite” is used in preference to “hermaphroditic.” Harlan Lewis (1962) will certainly be astounded to learn that his “catastrophic selection as a factor in speciation” (p. 364) has been transformed into “catastrophe speciation.” How do catastrophes speciate, anyway? A final comment is necessary regarding the illustrations in the book. Some are excellent, especially the ones provided by M. C. F. Proctor, but others actually detract from the overall quality of the book. Many of the black-and-white photographs, such as those of purple-and-white-flowered fritillaries on p. 196 or violets on p. 365, do not show what the author intended and could be eliminated to good effect. As stated in the preface to the book that I edited for Chapman and Hall (Wyatt, 1992), the field of plant reproductive biology has grown vigorously in the past 20 years and has become increasingly heterogeneous. Evidence of this can be seen by a quick perusal of any recent issue of American Journal of Botany, whose pages over the past several years have been dominated by exciting new papers in this field. The task of summarizing all of this progress in a thorough and evenhanded way is now Herculean. Richards has made a valiant effort to collect in one place a diverse assortment of otherwise isolated studies. I applaud his effort, which should lead to easier identification of new problems for investigation. Here I will comment on some of the developments in plant reproductive biology that I see as important and which I feel Richards has not given adequate coverage in his second edition. First, I believe that we are only beginning to realize the potential value of the rapid growth in our knowledge of plant phylogenies. Phylogenetic reconstructions at all taxonomic levels, using new molecular data as well as reinterpreted morphological data, provide insights into when and where reproductive innovations arise. In this way we can dissect out the relevant character transformations to see if they are associated, for example, with a shift in habitat or ecological associates. By mapping reproductive traits onto a phylogenetic tree constructed from independent data, we can determine how many times that trait has evolved, gaining an understanding of the selective forces involved and of how likely it is that parallel evolution can occur. In this regard, the recent work of Armbruster (1997) and Johnson, Linder, and Steiner (1998) is exemplary. David Lloyd pioneered the application of general selection models to the evolution of plant reproduction. He and his coworkers have influenced others to use this approach to great effect. Often the results of these models have helped to guide our thinking in general about these issues and to direct our attention to the most important parameters to measure. Even though they are not necessarily accurate reflections of the real world, they offer insights into how plants function and the likely magnitude of differences that character changes will make. Following in Lloyd's tradition, Morgan and Schoen (1997) illustrate the utility of this approach. It is unfortunate that Richards's book fails to capture the essence of what this approach has to offer. Another approach that has prospered in plant reproductive biology, but which seems to be downplayed by Richards, is interpretation of reproductive traits in ecological terms. The original view that all transitions in breeding systems should be driven by genetic consequences has been expanded to accommodate a wider range of driving forces. For example, herkogamy and dichogamy, originally interpreted strictly as “outcrossing mechanisms,” may more often have been shaped by selection to prevent interference between the male and female functions of hermaphroditic flowers (Lloyd and Webb, 1986). The Darlingtonian hegemony has been broken, without the necessity of rejecting all of its principles. But you would never realize this from a reading of Richards's book. And certainly, I agree with Richards's contention in the preface (p. x) that “evolutionary genetics still underpin all plant breeding system work, but in almost no case is any genetics undertaken.” It appears that molecular genetics has eclipsed Mendelian genetics, often to the detriment of those of us most interested in the ecology and evolution of plant reproduction. Useful information has been contributed (e.g., about the existence and structure of the S-locus in self-incompatible plants), but other basic mysteries that could be explained by classical genetic approaches have been shunted aside (e.g., if “late-acting self-incompatibility” is indeed under genetic control). We often assume a genetic basis to a trait of interest and further assume that genetic variation exists on which selection can act. Personally, I am disappointed at what quantitative genetics has contributed thus far to our understanding of the evolution of plant reproductive characters. The difficult and time-consuming experiments and the complex and ambiguous statistical designs are daunting, and the results thus obtained seem so hard to interpret and apply back to the natural situation that little seems to be gained for the effort invested. Perhaps Ruth Shaw and others will soon show us a way out of this conundrum. Three other technical approaches that have provided valuable new information also appear to me to be reaching the limits of their utility. The ability to use genetic markers in conjunction with statistical estimators to quantify outcrossing rates was a major breakthrough in the early 1980s. It enabled us to compare plant populations or species in terms of what was actually achieved with respect to self- vs. cross-pollination and provided other insights into mating within populations. Attempts to extend this approach to a finer scale, to examine family-level outcrossing rates, have been problematical, even in cases where inferences are not limited by sufficient genetic polymorphism. It appears that newer, more robust statistical methods need to be developed. The concept of functional gender was envisioned by Lloyd (1980) as a way to avoid classifying plants into typological categories and, instead, to quantify the degree to which they contributed to the next generation as male vs. female parents. This approach provided useful insights into the evolution of dioecy from distyly, but the calculations are tedious for most cosexual plants, depending on data that are technically difficult to obtain. Thus, the approach, though conceptually brilliant, has had less impact empirically than one might have hoped. Nevertheless, it at least deserves mention by Richards. On the other hand, paternity analysis has provided useful tests of some of our pet ideas regarding gene flow and sexual selection in plants. Again, it is often fraught with analytical and interpretational problems, even when sufficient genetic polymorphism exists. Many of the statistical “solutions” to the lack of resolution, such as assigning “partial paternity,” are unsatisfying in the extreme. A few other arenas in which progress appears to be lagging include our understanding of female choice, male: female allocation, and pollinator effectiveness. Work on female choice seems not to have progressed beyond merely demonstrating in a range of plant species that progeny of certain crosses are favored on a given maternal plant. The mechanistic basis of this phenomenon remains unexplained. Models that predict mating-system evolution often depend critically on measurement of reproductive investment in male vs. female function. Yet we seem to have made little progress in devising meaningful ways to quantify these parameters. Many recent authors have discovered that plant-pollinator interactions seldom lead to highly coevolved and specialized relationships (e.g., Waser et al., 1996; Waser, 1998), in strong contrast to earlier ideas on the subject. Given that most plant species are generalists, we need better means of measuring pollinator effectiveness. The older tradition of merely listing all plant visitors and guessing which are most effective needs to be replaced with a more direct, quantitative assessment. We may then discover that features of certain flowers have been misinterpreted under a too-ready acceptance of the dogma of pollination syndromes." @default.
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