IS A BRYOPHYTE SPOROPHYTE A PARASITE? I was reading Chapter 2-1 of your book on Bryophyte Ecology; at the end, the 'Summary' states that the gametophyte supports a parasitic sprophyte. I believe this is a mistake that has caused long discussions among teachers in our introductory courses. The growing sporophyte causes no damage to the gametophyte and once in the open, it is photosynthetic. Clearly, there is a misconception of the relationship between the gametophyte and the sporophyte. To me, it would be like saying that my fingers or hands are parasitic on my trunk! What do you think? Claudio Delgadillo In reading my Chapter 2-1 Summary, Claudio Delgadillo has raised the question of whether we can truly consider the sporophyte of a bryophyte to be a parasite on the gametophyte. He argues that it would be like considering my hand to be a parasite on my body. He further argues that the sporophyte causes no damage to the gametophyte and that it is photosynthetic for part of its existence. Claudio's points are well founded, so I decided to explore it a bit on the internet to see how others consider the relationship. The first definition I found for parasite is: an animal or plant that lives in or on a host (another animal or plant); it obtains nourishment from the host without benefiting or killing the host. I agree that the sporophyte is capable of fixing photosynthate, but it also derives nourishment from the gametophyte, so I feel that part of the definition is satisfied. Thus, the question becomes, is the sporophyte a different plant by virtue of being a different generation? To follow the human analogy, is a fetus a parasite on its mother? I would consider that the sporophyte does do the gametophyte harm, as discussed in literature cited in Chapter 5-9 of my bryophyte ecology book <http://www.bryoecol.mtu.edu/>. Its requirement for energy from the gametophyte greatly reduces the biomass gain of the gametophyte compared to those gametophytes not supporting a sporophyte, at least in the species studied. At this point, it would seem one could argue either way. Convention becomes our guide, as language is a product of human use. The University of British Columbia web site on bryophytes states "The sporophyte grows partly as a parasite on the gametophyte." The University of Illinois Chicago website states "Sporophyte is a parasite." The World of Biology Website states "In Seed Plants, the Gametophyte is a very small Parasite of the Sporophyte." "The young sporophyte, as for ferns, develops initially as a parasite on the gametophyte." Similarly, in Brittanica, "Among the gymnosperms the male gametophyte is much reduced and is a parasite on the sporophyte for only a short time." The Berkeley Tree of Life website refers to algae: "Small sporophyte is dependent on a larger dominant gametophyte." Clearly the concept of a plant being parasitic on its other generation is not unique to bryophytes. On the other hand, a German website, referring to bryophytes, states "If these both wouldn't belong to the same species, the sporophyte could be called a parasite on the gametophyte." This brings us full circle. Can we legitimately call a species a parasite on itself? It seems to me we have broadened our use of the term to include such instances. We have a history of doing this, particularly in ecology, where usage of the term plant strategy became very controversial, but it has stuck in the literature. Janice Glime ++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++ RESPONSES I would vote no - the sporophyte is not a parasite on the gametophyte. Why no- because it is part of the life cycle of the system, and the gametophyte clearly benefits from the interaction (offspring production) . Although I suspect that the connected gametophytic tissue dies in the process. Clearly we can ask what are the true effects of the interactions to both players (I am ignoring the male/sperm and spores) I suspect we will need to carefully argue some of the possible +, - or 0 effects. Using parasite sounds provocative and draws interest from others (etc the public/students). To think a fetus / seed is a parasite on the parent is another way to look at parent/offspring interactions and can even be very useful in tapping into theory from another field to understand this relationship/interaction. Another interesting case are the males of some deep sea fish that are very small and are parasitic on females :) . (these are clearly different individuals) I guess diversity does interesting things to definitions take care Nicholas -D. Nicholas Mc Letchie Ph 859 257 6786 fax 859 257 1717 http://www.as.uky.edu/biology/faculty/mcletchie/ Department of Biology 101 Morgan Building University of Kentucky Lexington, KY 40506 +++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++ Janice, I say yes. It is definitely not the same as saying my hand is parasitic on the rest of my body. I do not believe that there is a similar situation in animals for a comparison. Sincerely, Keith Bowman ++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++ No. If we say yes, I think that we're abusing the word "parasite''. I agree with what the German website says, since both the gametophyte and sporophyte belong to the same species, we can't call the sporophyte a parasite. As to the definition "parasite-" it obtains nourishment from the host without benefiting or killing the host'', I'll say that the sporophyte does benefit the host. Without a successful reproduction system, there will be no more gametophytes. That's how life is. Hua -++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++ This is a very interesting question which brought back many memories of my first academic job interview at the University of Florida. My interview seminar dealt with carbohydrate metabolism in source and sink tissues. While the main part of my talk dealt with carbohydrate metabolism in potatoes, I had recently done work on carbohydrate metabolism in the gametophyte and sporophyte of several moss species. I was asked the question, "If mosses lack vascular tissue how could you talk about a source and a sink." I know this is a little off your question but it got me to thinking about the relationship between the sporophyte and gametophyte. I don't think I would class the generations as being parasitic since each generation derives a benefit from the other. I think the association would be more mutualistic than parasitic. The sporophyte generation would get the more immediate benefit from the photosynthate from the gametophyte but, the moss as a whole would benefit due t! o genetic Cyndy Galloway ++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++ No. How about "matrotrophophyte" (noun) or "matrotrophophytic" (adj.) from matrotrophy. Matrotophy has common usage in zoology and botany. Why give the sporophyte the negative connotation as being a parasite? I see no value in this other than to convey the concept of partial dependence on the gametophyte. It is not a parasite anymore than a human fetus as you point out. Thank you for the opportunity to offer an opinion. Paul G. Davison UNA Box 5232 Department of Biology University of North Alabama Florence, AL 35632 Phone (256) 765-4434 Fax (256) 765-4430 http://www2.una.edu/pdavis/ ++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++ From my point of view one generation of a species can not be the parasite of the other generation. Without the sporophyte there would be not sexual reproduction of mosses. As a consequence, this would harm the evolution of mosses. So, there are two more biological arguments against this concept: 1. Mosses benefit from the alteration of generations in terms of evolution and dispersal. 2. Parasitism is an option (a thread) to the host without benefit for the host. Alteration of generations is far more than an option. Warm regards, Ralf. -Ralf Reski Professor Dean Faculty of Biology Head Plant Biotechnology University of Freiburg Schaenzlestrasse 1 D-79104 Freiburg Germany Fon: +49 761 203-6969 Fax: +49 761 203-6967 ralf.reski@biologie.uni-freiburg.de http://www.plant-biotech.net/ +++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++ Yes, but I'm open to alternate language and look forward to the responses you get! A fetus hormonally mines its mother of calcium and nitrogen, so to me it is a very well-adapted parasiste...even using specialized globins for oxygen transfer. FWIW: I never use "fertilization" when I mean "syngamy" because of the obvious inaccuracy and confusing interpretation of the former...for botanists "fertilization" often means application of minerals, compost, etc. rather than union of gametes...and females are fertile whether male gametes are available or not...so just as NASA forced physiologists to stop using geotropism when they mean gravitropism...we do need to make sure our terms accurate. An interesting thread! ross +++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++ They are separate organisms, despite their genetic relationship. -__________________________________ Martha E. Cook Associate Professor of Botany Illinois State University Department of Biological Sciences Campus Box 4120 Normal, IL 61790-4120 USA Phone: 309-438-8549 Fax: 309-438-3722 E-mail: mecook1@ilstu.edu http://www.bio.ilstu.edu/cook/ +++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++ Parasitism is a type of relationship that involves two individuals of different species (from Wikipedia: “two organisms which are phylogenetically unrelated”). Thus, I think it is not well used to describe the relationship between gametophyte and sporophyte in bryophytes, because in this case there is one only species (and maybe one only individual, or how many individuals are there in a fructified gametophore?). As an exception, parasitism can occur within the same species, as in the case of brood parasitism: brood parasites lay their eggs in the nests of hosts of the same or different species. Intraspecific brood parasitism has been reported for a range of insect, bird and fish species. In addition, in parasitism one individual (the parasit) benefits and the other (the host) is harmed. In bryophytes, gametophyte growth may be negatively affected by sporophyte production, but spore formation may contribute to genetic diversity and thus to environmental adaptability and ecological success. Thus, it is debatable if the supposed “host” (gametophyte) is harmed or benefited or both. I think that the term “parasitism” is being successful because of a simple question of fashion, in the line of similar “sensationalist” terms such as “plant strategies” or “invasive alien plants”. There is a term (auxotrophy) that sometimes has been used to describe the relationship between gametophyte and sporophyte in bryophytes. From Wikipedia: “Auxotrophy is the inability of an organism to synthesize a particular organic compound required for its growth (as defined by IUPAC). An auxotroph is an organism that displays this characteristic; auxotrophic is the corresponding adjective. Auxotrophy is the opposite of prototrophy. In genetics, a strain is said to be auxotrophic if it carries a mutation that renders it unable to synthesize an essential compound. For example a yeast mutant in which a gene of the uracil synthesis pathway is inactivated is a uracil auxotroph. Such a strain is unable to synthesize uracil and will only be able to grow if uracil can be taken up from the environment. This is the opposite of a uracil prototroph, or in this case a wild-type strain, which can still grow in the absence of uracil. Auxotrophic genetic markers are often used in molecular genetics. Researchers have used strains of E. coli auxotrophic for specific amino acids to introduce non-natural amino acid analogues into proteins. For instance cells auxotrophic for the amino acid phenylalanine can be grown in media supplemented with an analogue such as para-azido phenylalanine. Aminoacyl tRNA synthetases recogonize the analogue and catalyze its binding to a tRNA which subsequently transfers the amino acid (non natural in this case) to a growing polypeptide chain during protein translation. It is important to remember that many living things, including humans, are auxotrophic for large classes of compounds required for growth and must obtain these compounds through diet (see vitamin, essential amino acid, essential fatty acid).” The term “auxotrophy” does not fit exactly with the relationship between gametophyte and sporophyte in bryophytes, but it may help. Probably this relationship is so unusual in nature that we need to invent a new term. With best regards, Javier -********************************** Dr. Javier Martinez-Abaigar Universidad de La Rioja Complejo Cientifico-Tecnologico Avda. Madre de Dios 51 26006 Logroño (La Rioja) (España, Spain) Tel.: 34+941+299754 Fax: 34+941+299721 javier.martinez@unirioja.es http://www.unirioja.es/ecophys/ ********************************* +++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++ Yes, I consider the sporophyte parasitic on the gametophyte. Matritrophy is key to sporophyte survival, but the word is too loaded. The sporophyte is dependent, like a child is on its mother, for as a child can breath so too may a sporophyte photosythesise (perhaps even using atmospheric carbon via its stomata, in the case of mosses). C. C. C. Chater, UK ++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++ This brings to mind that I once read something about excised hornwort sporophytes growing in vitro--can't remember details or the source. I once found a population of hornworts where rain runoff had completely covered the gametophytes, but the sporophytes (many of them several cm in length) appeared quite healthy and growing vigorously. There were many dozens of them, growing densely enough that the population resembled a patch of strange graminoids. The gametophytes I retrieved from the sandy soil were black and shriveled so far as I could see with a handlens. I did not examine any of them in the laboratory, but it did seem that the gametophytes were serving as little more than perhaps a sort of wick between sporophyte and soil solution. I am wondering if there have been any observations of further maturation in the relatively large chlorophyllose young sporophytes of certain mosses if they are excised and maintained in vitro? Dan Marsh ++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++ About the question, I think that the sporophyte is not parasite of gametophyte, first is the same species although a different generation. The development of sporophyte is necessary to complete the life cycle, so if the sporophyte complete its life and produce a lot of spores it will be good for the species ( including gametophyte). Is the root of a tree a parasite of the photosynthetic part of the tree? or Is the female gametophyte of angiosperms parasitic of sporophyte ? I think no. sincerely Felisa M. Felisa Puche Pinazo Departamento de Botanica Facultad de Ciencias Biologicas tel. 96 354 4633 Universitat de Valencia Dr.Moliner n 50, 46100- Burjassot Valencia. Spain +++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++ Thanks you very much of the interesting discussion on the use of the term "Parasite". I personally agree that the sporophyte is, at least partially, a parasite of the gametophyte. In human body, hands are not parasites as they are part of the body, whereas in plants, gametophytes and sporophytes are completely different entities. As one is haploid other is diploid, the genetic make up is at least partly different. Consequently, I would consider animal fetus a parasite on its mother. I think that is a evolutionary strategy to nurse the next generation. Animals that hunt and eat its own species are still consider carnivores (although it is a special kind of carnivory call cannibalism), why can't we call a species a parasite when it get its nutrition from another individual of the same species? Unless someone wants to invent a new term for this special kind of parasitism of plants. Best wishes Boon-Chuan Ho +++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++ *German website, referring to bryophytes, states "If these both wouldn't belong to the same species, the sporophyte could be called a parasite on the gametophyte."* The question is a bit semantic and perhaps even philosophical, but I think the above statement is correct. How can a sporophyte be defined as a parasite of a gametophyte, if the sporophyte's whole existence only aims at producing more gametophytes like the one it is sitting on? True, the new gametophytes will not be genetically identical with the original one, but that is not the point here, particularly since the gametophytes of many if not most bryophytes are capable of producing clones of themselves if and when they "have to". I think the term parasite is wrongly applied here. Best regards, Johannes -Dr. Johannes Enroth PhD, University Lecturer, Bryologist Dept. of Biological and Environmental Sciences P.O. Box 65 FIN-00014 University of Helsinki Finland +++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++ Dependant vs parasitic? Is this distinction fouynded on whether the sporophyte actively enforces a regieme of growth and development onto the gametophyte, the result of which is the nutritional interface and protective structures produced by gametophyte. Does the gametophyte (as does any mother) actively facilitate the growth and development of the sporophyte? Does a chemical stimulus from the sporopyte initiating a signal transduction cascade (or whatever transmits that signal) in the sporophyte constitute the same kind of enforcement as a tick sticking its mouthparts into you? I would argue that the relationship between sporophyte and gametophyte stands in sharp contrast to the relationship between host and parasite, regardless of nutritional similarities in both types of relationship. regards Matt _______________________________________ Matt Renner PhD candidate The Wardle Lab School of Biological Sciences Heydon Laurence Building A08 University of Sydney, NSW 2006 (02)9351 5118 041 585 2205 ++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++ There is a whole generation of my high school students who learned a rhythmic "choral reading" of a short poem that went like this: In (pause) mosses, the sporo-phyte is a para-site on the fe-male gameto- phyte. I stood at the front and waved my arms around like a choir director as we said this over and over. They were very young high school sophomores, and I did whatever I could to keep them focussed and have a little fun along the way. They were also quite attentive after I told them that mosses and ferns can only do it in the rain....smile. Sadly, with the advent of standardized state tests and therefore, standardized curriculum topics, there is no longer time to teach a survey of either the plant or animal kingdoms. I guess when we retire we're allowed to reminisce about the good old days. Maggie Ray ++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++ Perhaps the term hemi-parasite may better sum up the situation in most, but not all, bryophytes. I think the situation in mosses may be more or less unique. In Marchantia, or other marchantioid hepatics, the sporophyte undergoes much of its development without the apparent need for photosynthesis. Riccia is perhaps the extreme case but even in Asterella, Marchantia, Lunularia, Reboulia, the carpocephalum contains photosynthetic tissue but it may be of use (to the sporophyte itself) only in the final stages where the sporophytes are elevated above the thallus. Jungermanniales do not, as far as I am aware, have photosynthetic tissue in their sporophyte. I wouldn't consider my arm, leg, head even, parasitic on the rest of me because each part derives all it needs (I think!) from the whole. If one looks at the case of Buxbaumia, what do we call the maturing sporophyte - which is very much larger and more conspicuous than the gametophyte (which, I hasten to add, I have never actually examined or seen). If one looks at Ephemeropsis, and perhaps to a slightly lesser extent Viridivellus, where the gametophyte is about the size of the sporophyte or may, because it is like a protonemal felt, be the dominant part, we have perhaps a 50:50 situation. In many instances the moss sporophyte is the lesser party. But, if we have to draw analogies because our minds like to have comparisons or places in which to put things, I think the instance of the animal foetus is probably the equivalent of the situation in a bryophyte - no, moss sporophyte. In hepatics, the sporophyte perhaps comes nearer to a truly parasitic relationship - but this then gets back to the philosophical argument about the possibility of being parasitic on oneself. The hornwort sporophyte is perhaps nearest to independent existence, but it depends on sperm (possibly foreign in the sense of being from another plant) being subsumed in nuclear fusion by the gametophyte egg cell to form the zygote. Perhaps not entirely unrelated to this discussion is the thesis once put forward that the chloroplast was originally a foreign organisms captured by another and subsumed in the dominant partner's make up. There is chloroplast DNA after all. Perhaps the molecular people can answer the question - how different is the chloroplast DNA to, nuclear DNA or ribosomal DNA? Regards, Rod Seppelt. Prof. Rod Seppelt, Australian Antarctic Division Channel Highway, Kingston, Tasmania 7050, Australia ph: +61 (03) 62 323 438 FAX: +61 (03) 62 323 449 email: rod.seppelt@aad.gov.au ++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++ Why not a hemi-parasite? I wouldn't consider my arm, leg, head even, parasitic on the rest of me because each part derives all it needs (I think!) from the whole. If one looks at the case of Buxbaumia, what do we call the maturing sporophyte - which is very much larger and more conspicuous than the gametophyte (which, I hasten to add, I have never actually examined or seen). If one looks at Ephemeropsis, and perhaps to a slightly lesser extent Viridivellus, where the gametophyte is about the size of the sporophyte or may, because it is like a protonemal felt, be the dominant part, we have perhaps a 50:50 situation. In many instances the moss sporophyte is the lesser party. But, if we have to draw analogies because our minds like to have comparisons or places in which to put things, I think the instance of the animal foetus is probably the equivalent of the situation in a bryophyte - no, moss - sporophyte. In hepatics, the sporophyte perhaps comes nearer to a truly parasitic relationship - but this then gets back to the philosophical argument about the possibility of being parasitic on oneself. The hornwort sporophyte is perhaps nearest to independent existence, but it depends on sperm (possibly foreign in the sense of being from another plant) being subsumed in nuclear fusion by the gametophyte egg cell to form the zygote. Prof. Rod Seppelt, Australian Antarctic Division Channel Highway, Kingston, Tasmania 7050, Australia ph: +61 (03) 62 323 438 FAX: +61 (03) 62 323 449 email: rod.seppelt@aad.gov.au ++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++ Parasitism? I would say: no. The problem of course lies in the definition of 'parasitism'. The one given here uses the 'plant or animal' (as if nothing else would exist, and I think here you can clearly see how traditional and conventional this approach is) as the unity. May be, we should think about the sporophyte and the gemetophyte as expressions of one and the same genome, and, in a sense, parts of a same 'organism'? Hans de Bruijn ++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++ Its an interesting question. I checked the mening of parasitism by wikipedia: http://en.wikipedia.org/wiki/Parasite I think for bryophytes this word can not be used because: it should be between phylogentically different individuals and the parasite must cause harm to the host: ("It is important to note that "benefit" and "harm" in the definition of parasitism apply to lineages, not individuals. Thus, if an organism becomes physically stronger as a result of infection but loses reproductive capabilities (as results from some flatworm infections of snails), that organism is harmed in an evolutionary sense and is thus parasitized.") Since the relationship between the gametophyte and sporophyte is a necessity for reproduction (well...hm. depending on if the species use sexual or vegetative reproduction) it is an evolutionary benefit. Perhaps its time to come up with a new term for the special relationship between gametophytes and sporophytes in bryophytes (a relationship that causes temporary harm to the host but is an evolutionary benefit in the longer term). But on the other hand: it is very similar to other cases as you already mentioned. Cases that are not considered to be parasitic. So, I do not know. Unfortunately, the world is usually much more complicated that systematist and language allows it to be. Annika Jagerbrand +++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++ Perhaps the term "hemiparasitic" could be better appropriate? René Schumacker ++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++ This is interesting but I think you have already answered the question with what you presented. Maybe I'll make a misstep here, so anyone correct me if I do. Since the gametophyte always gets the benefit of sexual reproduction from the sporophyte, and since the gametophyte produces the cells that lead to the creation of a sporophyte, I simply can't see how the term "parasitic" applies to a sporophyte. In essence, I don't see how one part of an organism is parasitic on another part. As mentioned before, a fetus (imperfect analogy since that is more like asking if a spore is parasitic on the gametophyte - both are the same generation, and still, as you well know the human placenta is not of the opposite generation from the mother so it is even a weaker example than a sporophytic placenta) is not a parasite and neither is a tumor. A tumor is also an imperfect analogy, but closer to a parasite in my mind in that there is no reproductive benefit to the host (at least in most cases). However, the tumor is NOT another organism and therefore, I would argue that it cannot be considered a parasite. I understand the desire to use parasitic because it is provocative, but I think the term "dependent" is appropriate and that we should be clear to make this distinction. If the concern is how to characterize sporophytes that can provide some photosynthate for themselves, I would use the term "semi-dependent". As for the the chloroplast being enslaved by the cell after endosymbiosis, one should realize that it is a two-way street benefiting both former organisms - a real mutualism. To partially answer Rod's question about differences in DNA, I believe the G:C content is quite different, but more fundamentally (in my mind) the chloroplast genome is a circuluar plasmid (just like bacteria) rather than a set of chromosomes (like eukaryotes) and the promoter structure is very different between nuclear and chloroplast genomes. I couldn't be more convinced that the chloroplast is essentially a bacterium within a eukaryote, but I won't go into all the various sorts of evidence. -Dave Hanson +++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++ I believe that the term parasite reflects the mode of life of a species in relation to a different species. This is a species fitness issue having population effects "In the population interaction called parasitism, one population (the parasite) benefits, and the other (the host) is harmed by the relationship." Because the plant sporophyte is just a different life phase of the same species as the gametophyte, their relationship is no more parasitic than that of a eutherian mammal's fetus. I both cases (and in the florideophycean red algae and cases of viviparous non-mammals) the fitness of the species is improved by maternal provision of nutrients to the young. I call this "plant matrotrophy," a term first used in discussions of viviparity in animals. The classic experiments of Adrian Browning and Brian Gunning demonstrated experimentally that matrotrophy improves fitness in Funaria hygrometrica, a result that can reasonably be extrapolated to all embryophytes (in the absence of evidence to the contrary). I would say that the plant sporophyte is matrotrophic on the gametophyte for at least some period of time during early embryo development. Linda Graham +++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++ It sounds like a tricky question if the sporophyte is parasitic or not, in bryophytes. I think Ros Seppelt gave a nice introdution to the differences and problems involved, thank you. In any case, I also would exclude parts of the same organisms (and same generation) in the definition of parasitic. Regarding chloroplast, generally three things are indicated as supporting an endosymbiotic origin of it, a) a double membrane, b) and more interestingly, a DNA which is of the prokaryotic type (no istones), and, c) ribosmes of the bacterial type (different from those of eukaryotic cells), even more interestingly, the same reasons apply to mitochondria too. -Dr Carmine Colacino Lab. di Briologia - Dip. di Biologia DBAF Universita' della Basilicata Viale dell'Ateneo Lucano, 10 Potenza, Italy Ph. +39-0971-206234; Fax +39-0971-206233 Lab.+39-0971/205346: Mob.+39-329-3178399 Alt. e-mail: colacino@bryology.eu http://www.unibas.it/utenti/colacino/ http://www.bryology.eu/ ++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++ The assertion of a parasitic sporophyte assumes that there is no benefit for the gametophyte from the sporophyte. If one is to accept one of the basic tenets of evolution...the primary goal is the survival of the DNA into the future..., then the question seems to answer itself. The sporophyte is preserving the genetic code of the gametophyte by producing spores. The spores can survive inclement conditions, or even be transported in ways the gametophyte cannot. Obviously, bryophytes, with their infinite variety of asexual reproductive strategies, do not rely on sexually produced offspring. However, the fertile gametophyte spends a great deal of energy creating this "parasite". Extending the human metaphor, the "parasite" child is not only an energy drain in utero, but also for the first 15-20 years of life! This is an awesome investment of energy. It is obvious that the species, and thus the DNA would perish without offspring, and is thus clearly beneficial to the parent. So parasite is at best an inaccurate word for this relationship. Partner seems a better place to start if one is convinced that the two generations constitute a separate being. Ken Kellman 9870 Brookside Ave Ben Lomond, CA 95005 kkellman@sbcglobal.net (831) 336-8548 ++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++ On "parasitic", I tend to think that this is a bit harsh to apply to moss sporophytes, since the sporophyte represents offspring, and the gametophyte allocates nutrients toward their maturation, actively mobilizing their transfer as I'm sure you are aware through an elaborate system of coevolved membranes at the sporophyte/gametophyte juncture. When stressed, I'm finding that the maternal gametophyte jettisons the sporophyte to preserve herself, and this is to be expected in perennial species, which then live to reproduce another set of offspring hopefully. Clearly, the two generations signal each other, and the fitness of one is intertwined with the fitness of the other. I'm reminded of the stimulation of postfertilization perichaetial leaves and the function they may confer in protecting the young embryos of many pleurocarpous mosses (a phenomenon not expected in a parasitic situation). As for an alternative term, would your suggestion of "hemiparasitic" work? To me the analogy is more to a fruit on a tree. Maybe anything dealing with "parasitic" really connotes too negative for what is going on, however at the moment I have nothing to put forward, perhaps something will come to me. When I think about the generations and watch them grow, I tend to think of a single entity with two organs, rather than two separate entities vying for survival. Lloyd Stark +++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++++ Yes, I think that you have to consider the sporophyte as a parasite on the gametophyte. Considering Claudio's points I would say that the situation is different to the hand and the body because the sporophyte is genetically distinct from the gametophyte. Also, nobody doubts that mistletoe (Viscum album) is parasitic on the host tree, but it appears to do the tree little or no damage and does also photosynthesise. I think that what makes us feel uncomfortable is that the sporophyte is of some benefit to the gametophyte in that it does disperse its genes. Perhaps we could say that the sporophyte is a physiological parasite but a genetic symbiote, or is that too much of a mouthful? Jonathan Sleath ++++++++++++++++++++++++++++++++++++++++++++++++++++++ I shouldn't put the idea into university heads, but as an undergraduate at Queen's University in Kingston, Ontario in the 1960's I had to undergo a final oral (as well as written) examination. So the question of whether or not the sporophyte is parasitic would be a wonderful question for a student. It is entirely open-ended and would certainly tell if the student could think rationally and quickly on his or her feet. One question I was asked during my oral was whether or not viruses were alive. Nasty stuff. About the sporophyte - I'll leave that debate to those more knowledgeable than I! Allan Aubin