Concerning the Origin of Malignant Tumours by Theodor Boveri. Translated and annotated by Henry Harris

In the year 1902, I tacked onto the results of my experiments on the development of doubly fertilised sea urchin eggs the speculation that malignant tumours might be the consequence of a certain abnormal chromosome constitution, which in some circumstances can be generated by multipolar mitoses (Boveri, 1902). I had intended, even at that time, to put that assumption on a firmer footing in a separate article. But the scepticism with which my ideas were met when I discussed them with investigators who act as judges in this area induced me to abandon the project. I had to admit that a hypothesis in this field can be of value only if it leads to targeted new research and, above all, to new experimental investigations. And who ought it to be to decide to undertake such investigations if not the originator of the hypothesis himself? So, for some time now, I have carried out experiments that seemed to me to be relevant, admittedly without success so far, but my convictions on this score have nonetheless not been shattered.It was the appearance of an article by O. Aichel (Aichel, 1911) that first prompted me to take the decision to communicate the hypothesis, and the evidence that supported it, in greater detail, even though I had essentially nothing more to offer than I had done ten years previously. To explain the origin of tumours, Aichel's article relies on facts that I had established in sea urchins and that gave rise to my own conception of tumours. But the hypothesis that multipolar mitoses might generate tumours is amalgamated by Aichel with his own view, published earlier, that the initial event in the generation of a malignant tumour is the fusion of a tissue cell with a leucocyte(1). This notion casts the key elements of the hypothesis for which I am responsible in an essentially different light, so much so that it has become a matter of importance to me, now that my views have again become the subject of discussion, to set them out in their original form for readers familiar with the subject. But, in the end, I had another motive. The idea that might be a abnormal mitoses and malignant tumours has but it has been so that in the it is if at The that the idea to But can nonetheless and the more that now have might a of their with malignant tumours. might as a to that idea has been has been that some of the of malignant such as might be on tissue by their fusion in with of tumour with has not been but fusion with and is that malignant tumours that have been by the fusion in of the tumour with tissue of the but this has in the of the tumour is not at of and tumour have been and to in the so more than the tumour and to be responsible for the of the tumour this as a I have in of the of tumour it is that I am of in the that I the of facts and that I not set by But this article even more as is so the when an an a field with which is who has this to can to offer to investigators who have years and even of and to the tumour But is that has to be The tumour is a cell and, at the it is not that a who to certain in the cell might be to that the of tumours but that nonetheless their I that be with in some who this and in the that the tumour is a not a this that the the of a malignant tumour is to be not in the of the malignant but in the of the of the tissue in which the tumour not a is of his as an the that have met with a of the is that his views on the origin of malignant tumours by his on sea urchins and to my in the I not it as on me to my views of my it be without is only I and that is who and on put ideas that to my his views on the of abnormal chromosome in the of malignant tumours have been as as I can I have to might at the for their such in conception and in ideas and my own that the hypothesis I am now might elements that in as is as the originator of as a The chromosome that on in of the of but the of the and the of who had the of abnormal in tumours, and to have been the first to that the abnormal chromosome by mitoses was an of malignant this as a of the of tumour I only that in this field might not this I to my to my in in in and to the of the of of for their on the of tumours to me to be supported by the The of even the malignant tumours can be tissue The of this abnormal to be in the tumour not in their and malignant tumours have in I have to with who a the In my view, be a a tumour that in the as the tissue which it is and that to me that the of a tumour a malignant so is a of the as the appearance of a malignant tumour at some in tissue is a of with to the views discussed in that that not an event the but of tissue to as for of have a but have not in not for the of malignant tumours. in the of and by the of that which is a at the of the is by that the of the another for the of tumour own on which is discussed in some leads to that the of a malignant tumour is a is that when a tumour malignant that is again by a event the tumour and not by of a of not on the that malignant tumours might that in their But such to me to be with in which the more of malignant is a by the of that were at an is not that and cell a notion that the of even is a the that a malignant tumour might out of a cell that is for some and the idea that by the of some might a of elements of my of be as malignant tumour cell is a cell with a it has that a tissue cell In this I am in with the to which has the cell in this to and it is that this might for to without is a of has a of out of a not have and without has that if the in this had been the of generated this have the of the by a of this his to the a of cell not the of a new The of the more than a the of was on the assumption that the of a new that in a This was by the of tumour the assumption was that the by new the a new that be to in a views of with that was to a of evidence in of had to the in of cell fusion as a for the of of this it be that that to the malignant was to be a and not a in the and a of not the of a new The of that can the of malignant tumours to the of a sea of by a of tumour of at the of is that and not is the of a consequence of so and a without This idea to who the by for The key and so that on of their own in some has been some the origin of this to have been first set in in the in the but it was familiar to the is only at the when the of can be that and that that the The cell to be an and an in the that it not when the of the again that is not that in but that in is by the of the malignant this form of now the malignant cell as that has certain and to the of the this be to an cell to to and of to to the of But it is in the tissue of have that have to be can take this as it that elements in the cell in the malignant tumour The an of this of the idea even to the of tumour the of that a cell and that in malignant tumour in the rise to the can be a and the of such a and certain without the of the I have to such at a this I am to only by the of of the of can be of and The experiments that have been done to this in the that in of the of the as a at that the of the can be a so as a and eggs is a of much of an was in the field of The in it were on sea urchin experimental by and by with was on the of sea urchin eggs that his of chromosome the of their cell generation to the and the different of to to this In of the of is such that them generate of it is even in such the that out of the not abnormal is that the can generate only certain of And even with eggs that in this the which the is out to be rise to is to that tissue a a the to the cell so in it is the cell the to a in the cell by a of the of the experiments an different is not I set out the more the of the and the of the in and in that is of to in the in the of their only of the the of but in experimental it can be a that have a and of the and be in the can the that chromosome in the has in the the in the and The the and the amalgamated which now chromosome in this set is and a of the that a set is by of the by the of the fertilised In the the to But have to the of the chromosome that to the of that to as a that as the when the cell to of the of In this the of amalgamated at by the of the is only in the that the the set more a of the and the of were by in The by have been for in but in the of of the chromosome of the to the of the is only if the more take in the in the that to that the an and in an of The for this is that chromosome can only if only of the at the time can the of the nothing that to is another that be is a matter of which of the with a so that the not only have different chromosome but have different chromosome of that is by a for that a cell and and that it of the of the is in that it with chromosome chromosome and chromosome of of the The cell the chromosome as in In the out of the in of the the chromosome in that out of the chromosome a a of to the than a in an of of it that a cell this in some more than that and it that of the and some circumstances to certain in only cell a of of in cell a and in cell a is and, in cell is the the cell has a chromosome but nonetheless has This in and is a of abnormal of the to a is which can be in sea urchin for by of of the that But not with a set of but with a in the and in of the In this the that of the might a of of chromosome much to generate is to cell in This can be done with eggs and by them and In this the and of of to in the cell and to form the new as if cell had been this cell to of of and but the now rise to of a of the at the time, on the of and the that at of of chromosome be again much to in multipolar have a of that can be done without to the the of in which some in this can the that in with the the of such a in the of the To put the more can a with a of this is and, if it can it without to the first is as as even of be abnormal chromosome is by that mitoses The to the is that the of with by multipolar and that was that was not in the of a that the by as a that the that a and might this that multipolar mitoses have is a of an abnormal of I have carried out a of experiments with doubly fertilised sea urchin I gave a of this in and it in in (Boveri, my of this be to is the of the in This the of of chromosome of can be in sea urchin can them the the and the of of with a that the generate at a certain of the in to a of eggs of the without in a only a first to be in is that the of the by be responsible for the if it at be by the can the have be by the abnormal chromosome that the as a consequence of the multipolar experiments on and that eggs of eggs as as the chromosome can to explain the of is only assumption that it is the of that so for the the different such that only certain the cell to at it the and the the of which have and This at with a the in this of the the of the of a fertilised the the of a without eggs of the in view, it is that a at of the different of chromosome to of the the chromosome and only the of this which a eggs of this and more I the to my In the of in I the different of to abnormal with the that the I that in only a of the generate and the on is to be I to the the that for the of the to be some not the of the but only for the to form the to the of another that his to the be a the idea that rise to in that different have different to established in this for sea has by the experiments of certain different of sea was to that in the only the take in the development of the the This in a different that the the was a and an of a a of his and of this can be in the is that the set out in the on the of the sea urchin for have that of different to be in the mitoses of if the experimental to to a have with more and in the of the and in can the of have This assumption is by the that a chromosome of that have been in and and which with can be in some it as that have different in and it is this assumption that the of the tumour hypothesis I have put malignant tumour cell is and again I take the ideas of a cell with a abnormal chromosome this assumption be more some that might nonetheless be of the of in The by to the of to the chromosome in is and the is the in if the which has in the of the of a of such the chromosome in a And if experiments have that in some certain when in view, be in the chromosome The in which is is if that the of this is to the chromosome that as the as first by and in an essentially different by to an is generated when of the to and at the first cell of the In this of the have and have this a but is is a is was in In 1902, to the the of and that of the and this idea in greater a year To have at that time that which now were the chromosome in a was a evidence of of to the is evidence that of in with that only become in the of a of in the of the by the and of a a this it is but a to the that the of the cell on the of this is the that is supported by the experiments on doubly fertilised sea urchin experiments on sea urchins have me to that be that to and that to some is that for the of the cell and might the of the even but without it in that the of the cell to is as I in my might in be to of the cell and is that is a in cell that only cell to take when this is by some with if that were that cell their were cell tumour cell that without be generated if were In this the tumour cell the that were in the chromosome as the is that to cell that the on by the of to was in years when the of was by cell In it was that the of malignant was when were with but it when certain by the cell were the the hypothesis that that cell is with In this view, cell take when the of the of the is by some The of malignant tumour be to a of This of be of as a for that the for a is chromosome and for a it an chromosome such that can be by multipolar But it is that be a in the of the to a at this to the of but is not the idea that that cell the idea in his to this is that is to a in cell and is by experiments with that of time without I own is that the assumption that is a is than the idea that the chromosome of a tissue cell is so to the of the of the so as is in cell is it to me, when a in the has such an on the chromosome that it but a of this and cell take if a in the of the which be by the of And might not have to be This supported by the that tissue to in to different to me to be more than the assumption that This it to explain the of of malignant tumours. were a in the in cell at the on the the that is by the of different more than an abnormal can in the of might for of cell I that the of malignant tumours without to the that cell might that essentially the might in tissue if the of the of the of that the is The that when of tissue the as by to this the idea is that cell is the of if In the of tumour the is a of the to to the of this but the them the of such to the in the to this the and malignant tumours. The I have the in as I have that is a the of the of this not in the and malignant but in a that is to be The that the the in to an the of the I have malignant tumours to In of the hypothesis that that cell might the the of tumour in the idea that in tumours in malignant tumours the of in the of the to this idea is in of the of it is to the assumption that that act only as of cell the might that the in the of the of this not to take in is that is to an to a of this not the of it might explain the of that only in their to and generate tumours that tumours be by the in to an of have certain I have has a for the of tumours, but this in the I that the for cell is to a in the of the chromosome the be by an of not even the in but the abnormal of malignant tumours to the of I again the of the I have nonetheless be of the that the and the of cell In sea I was to in that an of to the of an to cell that cell a and another sea urchin in experiments with of eggs that have been can with more than of not the that the mitoses not but This a that rise to a cell this is that the of the is a in the cell in that in with that by the of the of the the of the cell by This the of to the tumour to me to be of some in the of this the of the in the had been to that the of cell was them that the to seemed to In the of and I have the of has been to that of that set the that a cell to But, in and the cell this the of the as I have a is not that of the that to cell the if this is it that a certain in the of the might a cell to again as as it is for it, as as such as not and the of the of the that is not by which of my more the of on the of the to mitoses can generate a of different chromosome so if hypothesis is in that a cell a tumour cell this is to the of certain in to an of of chromosome to the But, to with a some is only with an of with a the of the discussion, I again that the in my hypothesis is not the abnormal but a abnormal of the of it event that this generate a malignant tumour and can be a in certain by a of by of by that and to which I of the by is not the of but a that the of the I of as the in which a of is I to a more which not to be have much the as multipolar as the of has when the with of the not form an to the The that to only set of if of the chromosome has the to of the the cell of is that can to a of of chromosome to the if that is is of now as multipolar can take in tissue in The first is in the of the the that it has of the To by the of I have of this if at all, in not the of the certain to this might be to some abnormal for which is some that multipolar and, to be more mitoses can be is in the that I have that is by the of a cell that is In it is that cell is with in if at and new abnormal event to by in another is an in which is by abnormal of the and by of cell In the first a chromosome set is that is of the when cell is is that with the first of is a much that the of chromosome in the of has been out the origin and of tumours is as as my hypothesis is with my hypothesis, but not I not this but my to in my view, with my to the facts I in I have to it that if a malignant cell as that an this is in the not the this is the key of my hypothesis, it is now to the of this to the tumour more for to the I have put on the that I have discussed only that be that a by the of of the it has to be that of a that act might in the certain facts that the of tumours has such as the of by it that the in the tumour in not to the with which I have been I the origin of tumours that by I to out that it as as I am a that for at abnormal can be the assumption is that a tumour cell a has been in a not to and abnormal in to this But to the abnormal of and in by multipolar it to me, that the I in of importance in the origin of tumours. established that only on the have to that if the of certain malignant tumours that the of it is to that the notion of a of in the of malignant tumours, a is that have a in the and that have view, in of it, it was that the of was different in the The by the in that the was in and of the The on the of and was that in the tumour of the is in the so that to the malignant tumour that is is the of the But in the of that have to be which explain the the of the of this The has some but it has the that the of in is different that in the of is to the field of tumour research in the of have I first of matter that has not been as as I this matter me as of the as the of is different of tumours can in the of tumours has a that is in and This is to the a tumour might when the tumour is first now the of the tumours to have their the again be that to that had the abnormal is to in tumour has origin in a with is the of tumours. to be with tumours not as but out of But if to to a of the is the be the can that a of have been published in of the in an on the of in the published a of in which at that malignant tumours not but be this it might take to this I is and I am of that not take of origin is as I to it in to my hypothesis, a cell that a of as a consequence of an abnormal This is the of the for that the cell to so as to by But the abnormal that the tumour cell by the abnormal chromosome of the and be by the of this cell so as cell by the assumption that tumour is a cell is this an new on the by that malignant tumours at is I to this in another at this I to it the of the was the of the it be not the of the at in which to that it rise to an event that the that generate a malignant cell in only a of But the of an event such as the I am now be by abnormal of of the of to the of and it I have been that is a a of not admit the that might generate this not that the in the that to as a But is that is by the a of has that that is not a but a consequence of malignant has not been this in his to the of might that a more This is discussed in the that the of a of it can be as established that the of the tumour cell is by with is by the of the tissue of it that in different of the is the of different on the facts with The experiments on sea urchin eggs that I have me to that the in the different this is and in with hypothesis, tumour an abnormal of and an abnormal of can the of such is tumour cell that certain and has in in and in if at And it is if the of in at such the be but This explain the that malignant have on their of on their has now become a area of But it certain can the the can be for by the of the hypothesis not different of tumour can in is that for cell is abnormal of that the cell with the of that not that but not of the it is that a of different chromosome of which a on the In of hypothesis, this to the different malignant tumours that the tissue of origin to that to have the of the tissue of origin In the in which the for is the only only that cell The more the chromosome the the tumour the tissue of of that the in a sea of might be that not at not the that the chromosome might be in But to be is to the The was by in to his that new of be generated was of but at that time not to of years of results were to be in of chromosome and not offer an to The was by first on the in But the and of the of and his and in and is nothing in it that that was of the of and his at in and, to the on experiments with sea urchin of the of the chromosome not when who at time had with the in the of in the of the not that certain of chromosome if in the in in as can for when of cell a that only of a chromosome to of the and to this it is of that in it as if a in chromosome to chromosome in generate a