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KARL SAX
November2, 1892—October S. 1973
BY CARL P. SWANSON AND NORMAN H. GILES
As I view my contribution to the writing of our time, it seems to me to consist
of a double affirmative, saying first that an awareness and experience of Mature
is necessary to Man if he is to have his humanity, and saying in the second place
that that same awareness must have something of a re1tigious quality, the Italian
pieta, ifyo?~ will.
Mature is a part of our humanity, and without some awareness and experi-
ence of that divine mystery man ceases to loe man. When the Pleiades and the
wind in the grass are no longer a part of the human spirit, a part of the very
push at bone, man becomes, as it were, a kind of cosmic outlaw, having neither
fee completeness and the integrity of the animal nor the birthright of a true
humanity.
TH E S E W O R D S by Henry Beston from his now classic vol-
ume, Outermost House, strike us as uniquely applicable to
Karl Sax. He would not have been found wanting although
he, most certainly would have raised a quizzical eyebrow un-
less the term "religious quality" were stripped of any cloying
mysticism. He grew up and throughout his life remained
close to the soil, ant! he expressed in words ant! actions the
dignity, integrity, inner strength, and outer optimism that are
so often the legacy of such a birthright. He knew the wheat-
fields of southeastern Washington; he knew how to care for
and to harvest that which he had sown; and he knew the
wonder of growing things, whether these were plants or hu-
373
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BIOGRAPHICAL MEMOIRS
man beings. He acknowlecigect his birthright and was proud
of it, anct he shared it with those in both high ant] low places.
His public career was a long and ctistinguished one, but to
those who knew him privately as well, that record was but a
partial measure of a great and warm human being.
Karl Sax was born of pioneer parents in Spokane, Wash-
ington, on November 2, IS92; he stied in Meclia, Pennsylva-
nia, on October 8, 1973, less than a month shy of his eighty-
first birthday. His father, William L. Sax, was, at various
times, schoolteacher, county superintendent of schools,
farmer, businessman, ant! mayor of Colville, Washington. His
mother, Minnie A. Sax knee Morgan), was an artist and ama-
teur botanist. An exposure to plants and to the natural en-
vironment as well as the advantages of higher education were
very much a part of his early background. Sax entered Wash-
ington State College in 1912 to major in agriculture, and it
was here that he met Professor Ec~ward Gaines, a wheat
breeder in the Experiment Station. Gaines lect him into re-
search and uncloubtedly encouraged him to continue his
studies at the graduate level. As Sax once wrote, "Here ~
learned that one could have all of the pleasures of an agri-
cultural career without the financial headaches by going into
agricultural research work." This early experience with the
problems and techniques of plant breeding expanded into a
continuing ant} absorbing interest that was pursued through-
out his life. Other later studies brought him national ancT
international recognition, but they never fully replacect his
need to be close to the soil and to growing things.
Sax gracluated from Washington State College in 1916,
the year in which his first scientific paper appeared. Prior to
graduation he had married his cytology teacher, Dr. H ally
Jolivette. In the fall of 1916, she accepted an instructorship
at Wellesley College, and he entered the Bussey Institution
Graduate School of AppliecI Biology of Harvard University
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KARL SAX
375
to work under the direction of Professor E. M. East. He re-
ceivec] an M.S. clegree in 1917, but his graduate studies were
interrupter! by WorIct War I. He entered the army as a private
and was clischarged as a seconct lieutenant in the Coast Ar-
tillery in ~ 9 ~ 8.
Sax's first academic position was as an instructor in ge-
netics at the University of California, Berkeley, where he also
undertook cytological studies in the genus Crepis uncler Pro-
fessor E. B. Babcock. His stay in Berkeley was brief, however,
as was his next move to the private Riverbank Laboratories
in Geneva, Illinois, where he initiated his studies on wheat.
In 1920 he accepted a position at the Maine Agricultural
Experiment Station in Orono; here he completed his cloc-
toral thesis on wheat hybrids, and the D.Sc. was awarclect to
him in 1922 by the Bussey Institution. One of his colleagues
at Orono was John W. Gowen, anti they collaborated on a
number of occasions: the genetics, productivity, and root and
buc! selection of apples were their primary concerns. The
wheat studies were pursued almost as an avocation, but Sax
consiclered the papers dealing with wheat species and hybrids
to be his most important contribution cluring these early
years in large part because they were among the first of the
publisher! works that opened up what was then the new sci-
ence of cytotaxonomy in this country.
Sax remained at Orono until ~ 928 when he was appointed
associate professor of plant cytology at the Arnold Arbore-
tum ant! named to the faculty of the Bussey Institution Gracl-
uate School of Applied Biology, an affiliate of Harvard Uni-
versity concerned! with teaching ancl research in agriculture
ant! horticulture. Here he joinect a faculty of distinguished
biologists: W. M. Wheeler and C. T. Brues in entomology,
W. E. Castle and E. M. East in genetics, Oakes Ames in eco-
nomic botany, ant] I. W. Bailey in wooc! anatomy. The gracI-
uate student bocly must also have been a stimulating one be-
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BIOGRAPHICAL MEMOIRS
cause many of those who received their degrees from the
Bussey Institution were to become worthy successors to their
professors.
Sax was elevated to a full professorship in 1936, the same
year in which the dissolution of the Bussey graduate school
took place. (In the view of President Lowell, Harvard and
applied biology were incompatible.) The dissolution, how-
ever, necessitated a move of office, laboratory, and students
to the new Biological Laboratories in Cambridge; here he
taught courses in cytology and, for a while, genetics Sax
took over teaching the latter on the death of Professor East
in 1938. This move, on the other hand, did not terminate his
association with the Arboretum and the Bussey. His cytotax-
onomy studies continued, and many of his students lived in
the Bussey buildings during the summer months of their
graduate careers. For many of us this was during the latter
years of the Great Depression and under the lengthening
shadows of World War Il; to make ends meet we were en-
couraged by him to grow our own vegetables and to raid the
Arboretum for appropriate fruits.
With the retirement of E. D. Merrill, Sax was appointed
acting director of the Arnold Arboretum in 1946; in 1947 he
was named its third director. He held simultaneously the
rather empty title of superintendent of the Bussey Institu-
tion. But both administrative appointments were abruptly
terminated by Harvard University in 1954 as a result of his
vigorous but losing opposition to the proposal that the gen-
eral resources of the Arnold Arboretum books, herbarium
specimens, and funds be transferred to Cambridge as part
of a move for the consolidation of botany. Sax not only be-
~ .
lieved that the science of botany suffered when instruction at
the Bussey was terminated, and that it would deteriorate fur-
ther when interest in the Arboretum as a living center for
horticultural studies was lessened; he also considered the ac-
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KARL SAX
377
tion taken by the Harvard Corporation to be an outright
breach of trust. To combat the transfer, he enlisted the aid
of The Friends of the Arnold Arboretum and cooperated
with them when the group filed suit in Massachusetts against
the corporation. He contenclect that he, as director and as a
matter of principle, could not be party to the divestiture of
the Arboretum's resources without juclicial review and legal
approval. Sax remained as professor of biology in Cambridge
until his retirement in ~ 959, but the controversy left its mark.
The latter years were bitter ones: he was hurt by the aliena-
tion of some of his botanical colleagues and by the scientific
decline of the Arboretum that hac] been for so long a signif-
icant part of his productive years.
About thirty graduate students took their acivancecT de-
grees with Sax, and another fourteen spent their postdoc-
toral years in his laboratory. He is remembered ant! revered
with unabashed affection by these students; in his gruff but
quiet way he embraced them all ant! brought them into his
family. As he said, "My academic children seemed almost as
much a part of our family as our three sons."
.
Karl Sax established a solid! and enviable reputation both
in this country ant] abroad. He was as well known to nur-
serymen as to his fellow cytologists, ant! this was reflected in
his professional affiliations and in the honors bestowed on
him. He was a member of the Genetics Society of America,
serving as president in 1958; the Botanical Society of Amer-
ica he received its certificate of merit in 1956; American
Society of Horticultural Science; American Genetics Associa-
tion; Population Association of America; Planned Parent-
hooc! League, serving as president of the Massachusetts chap-
ter in ~958; American Academy of Social anct Political
Sciences; and the Radiation Research Society. He was electec!
to membership in the American Academy of Arts and Sci-
ences ~ ~ 94 ~ ~ anc! the National Academy of Sciences ~ ~ 94 ~ ),
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BIOGRAPHICAL MEMOIRS
to honorary membership in Phi Beta Kappa (1941) and the
Japanese Genetics Society (1956), and as foreign correspon-
dent to the French Academy of Agriculture (19461. The Jack-
son Dawson Memorial Medal of the Massachusetts Horticul-
tural Society was awarded to him in 1959, as was the Norman
I. Coleman Award of the American Association of Nursery-
men in 1961. He received an honorary doctoral degree from
the University of Massachusetts in 1965, from his alma mater
Washington State University in 1966, and from the University
of Maine in 1971. He was equally pleased, however, to be
named "Horticulturist of the Year" (1959) by the Student
Horticultural Club of the University of Massachusetts, and to
be grouped, by Katherine White in The New Yorker, with
Charles Sargent and Ernest "Chinese" Wilson as "a distin-
guished plantsman."
Sax was a national lecturer on the academic circuit for the
American Institute of Biological Sciences in 1957 and in 1962
for Sigma Xi. In 1951 he received the signal honor of being
asked to deliver the Lowell Lectures in Boston, choosing as
his topic world population problems.
The research and publication record of Karl Sax spanned
a period of fifty-five years (1916-1971) with but a brief in-
terruption for military service. The publications fall gener-
ally into three groups—horticulture, chromosomal studies,
ant! demography with considerable overlap of the first two
areas as much of the cytogenetic and cytotaxonomic work was
done on ornamental species in the Arnold Arboretum. The
horticultural aspects of Sax's professional career began with
his appointment to the Maine Experiment Station, where he
was much occupied with improvement of productivity in
apples. This interest, which initially involved propagation,
crossing, and sterility, was continued at the Arboretum, but
the focus of the work was now directed toward an under-
standing of the origin of the Pomoideae, the production of
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KARL SAX
379
desirable ornamental hybrids, and the means for dwarfing
well-known and useful varieties of nursery stocks. The dwarf-
ing of fruit trees hacl been practiced empirically for hun-
cireds of years before being introduced into the Americas but
the basis of dwarfing was not unclerstood in a scientific sense.
By experimenting with a wide variety of intervarietal, inter-
specific, and even intergeneric combinations of rootstocks
and scions; by the use of different interstocks between root
and scion; and by single ant! double bark inversions to block
the flow of nutrients through the phIoem, Sax contributed
significantly to an unclerstanding of the phenomenon, re-
~lucecl the variability of graft compatibility and growth, anti
simplified the techniques to the point where the average nur-
seryman could readily produce his own dwarfs.
In the area of plant breeding, Sax and his students- in
particular George Skirm were successful in creating a num-
ber of excellent hybrids that quickly found their way into the
ornamental tracle. He was especially proud of the graceful
cherry "Hally ~olivette," a hybrid between Prunus subbirtella
ant! P. apetela, which he namer] for his wife and frequent
collaborator. (The fact that Jolivette couIct be translated from
the French into "pretty little one" acicled icing to the cake of
tribute.) The magnolia "Dr. Merrill" honored his predecessor
as director of the Arboretum, while the crabapple hybrids
"Henry DuPont" and "Henrietta Crosby" were named after
two of the loyal Friends of the ArnoIct Arboretum, who were
also his personal friends ant! research sponsors. The
"Blanche Ames" honored a clistinguishec! botanical artist who
was also the wife of Professor Oakes Ames; the hybrid "Mary
Potter" was so named because one of the parent species was
Malus sargenti, named after her father, Charles Sargent.
Sax also producect a number of Forsythia hybrids. Beatrix
Farrancl, a well-known lanclscape architect, Friend of the Ar-
boretum, and designer of the gardens at Dumbarton Oaks
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BIOGRAPHICAL MEMOIRS
in Washington, D.C., was recognized by having a triploid hy-
brid named after her. This turned out to be a lesser improve-
ment than the tetraploid "Arnold Giant," winner of an award
of merit by the Royal Horticultural Society of England. The
"Arnold Dwarf" proved to be an interesting ground cover
but a meager producer of flowers. Still another of his Forsythia
hybrids, the "Karl Sax," was subsequently named by a nurs-
eryman who was testing it in his trial plots.
The chromosomal studies fell into two subcategories: cy-
totaxonomy and the effects of radiation and chemicals on
chromosome structure. As indicated earlier, his wheat studies
provided him with a doctoral thesis as well as helping to es-
tablish what was then the developing field of cytotaxonomy
in this country. He shared with the Japanese cytologists Ki-
hara and Sakamura the credit for discovering the role of
polyploidy and interspecific hybridization in the origin of
certain wheat species, a seminal work of great significance in
understanding the nature of some of our basic food plants.
Comparable studies, in which Hally Jolivette Sax often par-
ticipated, were carried out on a wide variety of groups grow-
ing or being tested in the Arboretum: Pomoideae, Pinaceae,
Rosaceae, CycIadales, Hamamalidaceae, Vitis, Yucca and
Agave, Rhododendron, Paeonia, Ulmus, and Platanus. The
karyotypes of Yucca and Agave were shown to be sufficient-
ly unique to cause them to be removed from the L.iliaceae,
and to be given familial status in the Agavaceae; moreover,
the complete fertility and regular meiotic pairing in the Lon-
don plane tree, a hybrid between Platanus occidentalis and P.
Oraentalis, demonstrated that separation by the Atlantic Ocean
for millions of years did not necessarily involve chromosomal
rearrangements and accompanying sterility.
It was Sax's interest in the American species of Tradescan-
tia, sparked no doubt by his collaboration with Edgar Ander-
son of the Missouri Botanic Garden, that led to the emer-
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KARL SAX
381
gence of radiation cytology out of what began as a cyto-
taxonomic survey of the Commelinaceae. Sax unclerstooc]
that chromosomal rearrangement must play some kind of
role in evolution and speciation and that the large size and
small number of Tradescantia chromosomes in the readily
available haploid microspores macle them ideal for experi-
mental purposes. Recognizing that X rays not only induced
mutations but chromosomal rearrangements as well, he ini-
tiatect his radiation studies in 1935. The atom bomb anct the
horrors of radiation exposure were nearly a clecacle in the
future.
Tradescantia paludosa was the species of choice, and the
following two clecades witnessed an extraordinary outpour-
ing of papers by Sax and his students papers that proviclect
qualitative and quantitative information on the frequency of
both inclucect and spontaneous aberrations. The implication
and transference of these data to problems of radiation ther-
apy, evolution, and speciation were inevitable, as was acicti-
tional information related to the effects of temperature, cell
cycle, (lose rate, and dose fractionation on the final frequency
of inducecl change. Sax was the father of radiation cytology,
anct he spawned a whole generation of "chromosome bust-
ers." In his later years, and particularly after retirement, Sax
turned to the chromosomal aspects of aging in seeds, and to
the radiomimetic effects of caffeine, insecticides, and chemi-
cal food acIditives.
While he vigorously pursued his horticultural and chro-
mosomal investigations, Sax still managed to take an interest
in and make a significant contribution to the area of clemog-
raphy. His initial entry into this field undoubtecITy stemmed
from his close association with his graduate mentor and now
colleague, Professor E. M. East; but it was probably fostered
as well by the interest of Castle ant! Brues in applied eugenics.
In 1923 East hac! publishecT Mankind at the Crossroads, a Mal-
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BIOGRAPHICAL MEMOIRS
thusian indictment of the present and future consequences
of-unchecked human fecundity in a world of limitect re-
sources and agricultural productivity. He acivocated a con-
scious and deliberate practice of birth control; in his words,
"parentage must not be haphazard." Sax was similarly Mal-
thusian. (Believing that what he acivocatec! publicly shouIct be
first practiced at home, he urged all of his graduate students
to read the so-callect "Bussey Bible," a collection of articles on
birth control.) The first of a continuing flow of articles ap-
pearec! in The Scientific Monthly in 1944, but the gist of his
thinking was set forth in his Lowell Lectures. The talks were
prepared for book form uncler the title Malthus and the Mod-
ern World; this was subsequently alterect to Standing Room
Only: The Challenge of Over-Population, which appeared in
1955 ant] was reissued in paperback in 1960.
Mil(l-mannered and retiring as he was in his personal re-
lations, Sax was actively aggressive in the Planned Parent-
hood League and in his demographic speeches and articles.
His local target was the restrictive birth control law of Mas-
sachusetts. These laws were subsequently changed by a ref-
erendum sponsored by the Planned Parenthood League—
but not before Sax hacl invoked the wrath of many religious
leaders and particularly those of the Roman Catholic Church
of Boston ant! its suburbs. (The Church proclaimed to its
flock that "birth control is against God's Law" ant! urger! all
parishioners to vote down the referenclum.) He viewer! the
harassment that resulted as a measure of the effectiveness of
his stanct, and so he continuer! his fight on a national scale-
believing, as has proven to be the case, that financial sect to
the undercleveloped countries without accompanying infor-
mation ant! aid regarding birth control was not only politi-
cally immoral but, in a human sense, ultimately self-clefeating
and cruel as well. He considered India a lost cause in this
respect, but he held high hopes for the Latin American coun-
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KARL SAX
387
With I. W. Gowen. The cause and permanence of size differences
in apple trees. Bull. Maine Agric. Exp. Stn., 310:1-8.
With I. W. Gowen. Permanence of tree performance in a clonal
variety and a critique of the theory of bud mutation. Genetics,
8:179-211.
The association of size differences with seed-coat pattern and pig-
mentation in Phaseolus vulgaris. Genetics, 8:552-60.
Bud and root selection in the propagation of the apple. Proc. Am.
Soc. Hortic. Sci., 20:244-50.
With I. W. Gowen. The place of stocks in the propagation of clonal
varieties of apples. Genetics, 8:458-65.
The relation between chromosome number, morphological char-
acters and rust resistance in segregates of partially sterile wheat
hybrids. Genetics, 8:301-21.
1924
The nature of size inheritance. Proc. Natl. Acad. Sci. USA,10:224-
27.
With E. F. Gaines. A genetic and cytological study of certain hy-
brids of wheat species. I. Agric. Res., 28: 1017-32.
With Hally Olivetti Sax. Chromosome behavior in a genus cross.
Genetics, 9:454-64.
The "probable error" in horticultural experiments. Proc. Am. Soc.
Hortic. Sci., 21:252-56.
Nursery stock investigations. Proc. Am. Soc. Hortic. Sci., 21.310-
12.
1925
Fertilization of apple orchards in Maine. Bull. Maine Agric. Exp.
Stn., 322:1-8.
1926
With Iva M. Burgess. Varieties of ensilage corn for Maine. Bull.
Maine Agric. Exp. Stn., 330:49-56.
Sweet-corn breeding experiments. Bull. Maine Agric. Exp. Stn.,
332:113-44.
Quantitative inheritance in Phaseolus. ]. Agric. Res., 33:349-54.
A genetical interpretation of ecological adaptation. Bot. Gaz.,
82:223-27.
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BIOGRAPHICAL MEMOIRS
Statistical methods in horticulture. Proc. Am. Soc. Hortic. Sci.
23: 141-49.
1928
Bud and root selection in the apple. Bull. Maine Agric. Exp. Stn.,
344:21-32.
Chromosome behavior in Triticum hybrids. (Verhandl. V. Internat.
Kongresses Vererbungs-wissenschaft, Berlin, 1927.) Z. Indukt.
Abstamm. Vererbungsl., suppl. 2: 1267-84.
1929
Chromosome counts in Vitis and related genera. Proc. Am. Soc.
Hortic. Sci., 26:32, 33.
Chromosome behavior in Sorbopyrus and Sorbaronia. Proc. Natl.
Acad. Sci. USA, 15:844, 845.
1930
Chromosome number and behavior in the genus Syringa. ]. Arnold
Arbor., 11:7 - 14.
With D. A. Kribs. Chromosomes and phylogeny in the Caprifoli-
aceae. I. Arnold Arbor., 1 1: 147-53.
Chromosome structure and the mechanism of crossing over. I. Ar-
noldArbor., 11:193-220.
Arnold Arboretum cytological laboratory report, 1929-1930. T.
Arnold Arbor., 11 :237, 238.
Chromosome stability in the genus Rhododendron. Am. I. Bot.,
17:247-51.
1931
The origin and relationships of the Pomoideae. I. Arnold Arbor.,
12:3-22.
Chromosome numbers in the ligneous Saxifragaceae. I. Arnold
Arbor., 12: 198-206.
Arnold Arboretum cytology laboratory report, 1930-1931. I. Ar-
nold Arbor., 12:299.
The smear technique in plant cytology. Stain Technol., 6: 117-22.
Chromosome ring formation in Rhoeo discolor Cytologia, 3:36-53.
Crossing over and mutation. Proc. Natl. Acad. Sci. USA, 17:601-3.
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KARL SAX
Plant hybrids. Arnold Arbor. Bull. Popul. Inf., 5:17-20.
1932
389
With E. C. Abbe. Chromosome numbers and the anatomy of the
secondary xylem in the Oleaceae. J. Arnold Arbor., 13:37-48.
The cytological mechanism of crossing over. }. Arnold Arbor.,
13: 180-212.
Chromosome relationships in the Pomoideae. I. Arnold Arbor.,
13:363-67.
Arnold Arboretum cytological laboratory report, 1931-1932. I. Ar-
nold Arbor., 13 :450, 451.
Meiosis and chiasma formation in Paeonia su~ruticosa. ]. Arnold
Arbor., 13:375 - 84.
The cytological mechanism for crossing over. In: Proceedings of the
Sixth International Genetics Congress, vol. 1, pp. 256-73. Brook-
lyn: Brooklyn Botanic Gardens.
Flowering habits of trees and shrubs. Arnold Arbor. Bull. Popul.
Inf., 6:14-16.
Review of Recent Advances in Cytology, by C. D. Darlington. Collect-
ing Net, 7:201 - 3.
1933
With Edgar Anderson. Segmental interchange in chromosomes of
Tradescantia. Genetics, 18:53-67.
With Hally Jolivette Sax. Chromosome number and morphology
in the Conifers. I. Arnold Arbor., 14:356-75.
With H. W. Edmonds. Development of the male gametophyte in
Tradescantia. Bot. Gaz., 95: 156 - 63.
Species hybrids in Platanus and Campus. J. Arnold Arbor., 14:274-
78.
Chromosome behavior in Calycanthus. ]. Arnold Arbor., 14:279-
81.
The origin of the Pomoideae. Proc. Am. Soc. Hortic. Sci., 30: 147-
50.
Chromosome numbers in Ulmus and related genera. J. Arnold Ar-
bor., 14:82-84.
With Susan Delano McKelvey. Taxonomic and cytological relation-
ships of Yucca and Agave. J. Arnold Arbor., 14:76-81.
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BIOGRAPHICAL MEMOIRS
1934
Interlocking as a "demonstration" of the occurrence of crossing
over. Am. Nat., 68:95, 96.
With Edgar Anderson. A cytological analysis of self-sterility in Tra-
descantia. Bot. Gaz., 95 :609-21.
With I. M. Beat. Chromosomes of the Cycadales. I. Arnold Arbor.,
15:255-58.
With Edgar Anderson. Interlocking of bivalent chromosomes in
Tradescantia. Genetics, 19:157-66.
With L. M. Humphrey. Structure of meiotic chromosomes in me
crosporogenesis of Tradescantia. Bot. Gaz., 96:353-62.
Cytology for students. (Review of Introduction to Cytology, by L. W.
Sharp.) Science, 80:407.
1935
.
With Edgar Anderson. Chromosome numbers in the Hamameli-
daceae and their phylogenetic significance. I. Arnold Arbor.,
16:210-15.
Chromosome structure in the meiotic chromosomes of Rhoeo dis-
color Hance. I. Arnold. Arbor., 16:216 -24.
The cytological analysis of species-hybrids. Bot. Rev., 1: 100-17.
Variation in chiasma frequencies in Secale, Vicia and Tradescantia.
Cytologia, 6:289-93.
The effect of temperature on nuclear differentiation in microspore
development. J. Arnold Arbor., 16:301-10.
With Hally Jolivette Sax. Chromosome structure and behavior in
mitosis and meiosis. I. Arnold Arbor., 16:423-39.
1936
The experimental production of polyploidy. i. Arnold Arbor.,
17: 153-59.
With Ladley Husted. Polarity and differentiation in microspore de-
velopment. Am. I. Bot., 23:606-9.
Polyploidy and geographic distribution in Spiraea. ]. Arnold Ar-
bor., 17:352-56.
Chromosome coiling in relation to meiosis and crossing over. Ge-
netics, 21:324-38.
With Edgar Anderson. A cytological monograph of the American
species of Tradescantia. Bot. Gaz., 97 :433-76.
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KARL SAX
1937
391
Eject of variations in temperature on nuclear and cell division in
Tradescantia. Am. I. Bot., 24:218-25.
Chromosome inversions in Paeonia suffruticosa. Cytologia, Fujii tu-
bilee Volume:108-14.
With Hally iolivette Sax. Stomata size and distribution in diploid
and polyploid plants. J. Arnold Arbor., 18: 164-72.
Chromosome behavior and nuclear development in Tradescantia.
Genetics, 22:523-33.
Review of Recent Advances in Cytology, by C. D. Darlington. I. He-
red., 28:217-19.
1938
The relation between stomata counts and chromosome number. I.
Arnold Arbor., 19:437-41.
Chromosome aberrations induced by X-rays. Genetics, 23:494-
516.
1939
The time factor in X-ray production of chromosome aberrations.
Proc. Natl. Acad. Sci. USA, 25:225-33.
With K. Mather. An X-ray analysis of progressive chromosome
splitting. J. Genet., 37:483-90.
With E. V. Enzmann. The effect of temperature on X-ray induced
chromosome aberrations. Proc. Natl. Acad. Sci. USA, 25:397-
405.
1940
An analysis of X-ray induced chromosomal aberrations in Trades-
cantia. Genetics, 25:41-68.
The effect of radiation on chromosome structure. Am. Philos. Soc.
Yearb., 1940:240, 241.
1941
With J. G. O'Mara. Mechanism of mitosis in pollen tubes. Bot. Gaz.
102:629-36.
With C. P. Swanson. Differential sensitivity of cells to X-rays. Am
J. Bot., 28:52-59.
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BIOGRAPHICAL MEMOIRS
The behavior of X-ray induced chromosomal aberrations in Allium
root tip cells. Genetics, 26:418-25.
Types and frequencies of chromosomal aberrations induced by X-
rays. Cold Spring Harbor Symp. Quant. Biol., 9:93-101.
1942
The distribution of X-ray induced chromosomal aberrations. Proc.
Natl. Acad. Sci. USA, 28:229-33.
The mechanisms of X-ray effects on cells. i. Gen. Physiol.,25:533-
37.
Diffusion of gene products. Proc. Natl. Acad. Sci. USA, 28:303-6.
1943
The effect of centrifuging upon the production of X-ray induced
chromosomal aberrations. Proc. Natl. Acad. Sci. USA, 29:18-
21.
With Robert T. Brumfield. The relation between X-ray dosage and
the frequency of chromosomal aberrations. Am. I. Bot.,
30:564-70.
1944
Population problems of a new world order. Sci. Mon., 58:66-71.
Soviet biology. Science, 99:298-99.
1945
The demographic dilemma. Science, 101 :325-26.
Lilac species hybrids. I. Arnold Arbor., 26:79-84.
Population problems. In: The Science of Man in the World Crisis, ed.
Ralph Linton, pp. 258-81. New York: Columbia University
Press.
1947
How new plants are made. Horticulture, 25(n.s.~: 127, 128.
Mechanism of heredity. Am. Fruit Grow., 67: 16, 28, 29.
Plant breeding at the Arnold Arboretum. Arnoldia, 7:9-12.
Temperature effects on X-ray induced chromosome aberrations.
Genetics, 32:75 -78.
Soviet science and political philosophy. Sci. Mon., 65:43-47.
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KARL SAX
393
The Arnold Arboretum during the fiscal year ended June 30,
1947. I. Arnold Arbor., 28:447 - 52.
The Arnold Arboretum of Harvard University. Arnold Arbor.
Bull., 10~31:9, 10, 24.
The Bussey Institution. Arnoldia, 7:13-16.
With Hally tolivette Sax. The cytogenetics of generic hybrids of
Sorbus. J. Arnold Arbor., 28: 137-40.
1948
The Arnold Arboretum during the fiscal year ended June 30,
1948. I. Arnold Arbor., 29 :422-28.
1949
The Arnold Arboretum during the fiscal year ended June 30,
1949. J. Arnold Arbor., 30:450-55.
John George Jack, 1861-1949. l. Arnold Arbor., 30:345-47.
The use of Males species for apple rootstocks. Proc. Am. Soc. Hor-
tic.Sci.,53:219-20.
. .
1950
Rootstocks for lilacs. Arnoldia, 10:57-60.
The cytological effects of low intensity radiation. Science, 112:332-
33.
Dwarf trees. Arnoldia, 10:73 -79.
Oakes Ames, 1874 - 1950. J. Arnold Arbor., 31:335 - 49.
The effect of X-rays on chromosome structure. I. Cell Comp. Phys-
iol., 35 (suppl. 1~:71-81.
The Arnold Arboretum during the fiscal year ended June 30,
1950. J. Arnold Arbor., 31:430-34.
The effect of the rootstock on the growth of seedling trees &
shrubs. Proc. Am. Soc. Hortic. Sci., 56:166-68.
Population and agriculture. In: Twentieth Century Economic Thought,
ed. Glen Hoover, pp. 647-68. New York: Philosophical Library.
1951
Biological resources as a factor in international understanding. Sci.
Mon., 72:300-305.
Can the earth feed its millions? UN World, 5:22-25.
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394
BIOGRAPHICAL MEMOIRS
Photosynthetic energy via agriculture. Proc. Am. Acad. Arts Sci.,
79:205-11.
The Arnold Arboretum during the fiscal year ended June 30,
1951. i. Arnold Arbor., 32:412-16.
Food resources and population growth. Bull. At. Sci., 7:105-7.
Population problems in world development. In: Social Progress
Through Technology: The Human Conditions of Economic Growth,
pp. 4-6. (A week-end conference in four panels.) MIT Foreign
Student Summer Project. Cambridge, Mass.: MIT Press.
1952
With Henry Luippold. The effect of fractional X-ray dosage on the
frequency of chromosome aberrations. Heredity, 6: 127-31.
With E. D. King and H. A. Schneiderman. The effects of CO and
O on the frequency of X-ray induced chromosome aberrations
in Tradescantia. Proc. Natl. Acad. Sci. USA, 38:34-43.
The Arnold Arboretum during the fiscal year ended June 30,
1952. i. Arnold Arbor., 33:403-9.
1953
Interstock effects in dwarfing fruit trees. Proc. Am. Soc. Hortic.
Sci., 62:201-4.
Enough for all? (Review of The Road to Abundance, by I. Rosin and
M. Eastman.) J. Hered., 44:203, 204.
The Arnold Arboretum during the fiscal year ended June 30,
1953. J. Arnold Arbor., 34:412-16.
Review: Symposium on chromosome breakage. Science, 118:658,
659.
With H. Kihara. Genetics in the U.S.S.R. I. Hered.,49~4~:132, 158.
1954
The control of tree growth by phloem blocks. I. Arnold Arbor.,
35:251-58.
Here's an easy way to dwarf trees. Better Fruit, 49:9, 10.
Population problems of Central America. Ceiba, 4: 153 -64.
Stock and scion relationship in graft incompatibility. Proc. Am. Soc.
Hortic. Sci., 64:156-58.
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KARL SAX
1955
395
With E. D. King. An X-ray analysis of chromosome duplication.
Proc. Natl. Acad. Sci. USA, 41:150 - 55.
The effect of ionizing radiation on plant growth. Am. I. Bot.,
42:360-64.
With E. D. King and H. Luippold. The effect of fractionated X-ray
dosage on the frequency of chromatic and chromosome aber-
rations. Radiat. Res., 2:171 - 79.
Evaluation of the recombination theory. I. Cell. Comp. Physiol.,
45(suppl. 2~:243-47.
Plant breeding at the Arnold Arboretum. Arnoldia, 15:5-12.
With A. G. Johnson. Induction of early flowering of ornamental
apple trees. J. Arnold Arbor., 36: 110 - 14.
Dwarf trees with bark inversion. Am. Fruit Grow., 75~3~:38, 39.
Standing Room Only: The Challenge of Overpopulation. Boston: Beacon
Press.
Pflanzenzuchtung im Arnold Arboretum. Dtsch. Baumsch.,7: 177-
83.
1956
What's new in plant propagation? Natl. Hortic. Mag., 35:116-18.
With Alan Q. Dickson. Phloem polarity in bark regeneration. I.
Arnold Arbor., 37: 173-79.
Paste the poison ivy. Arnoldia, 16:5-8.
The story behind dwarf fruits. Horticulture, 34(n.s.~:203, 233.
The population explosion, pp.3-61. Headline Series, Foreign Pol-
icy Association no. 120.
Review of Chromosome Botany, by C. D. Darlington. Science,
124:688.
1957
The control of vegetative growth and the induction of early fruit-
ing of apple trees. Proc. Am. Soc. Hortic. Sci., 69:68-74.
The effect of ionizing radiation on chromosomes. Q. Rev. Biol.,
32: 15-26.
Dwarf ornamental and fruit trees. Proc. Plant Propagators Soc.,
7: 146-55.
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396
BIOGRAPHICAL MEMOIRS
1958
The juvenile characters of trees and shrubs. Arnoldia, 18:1-6.
The genetic future of man. In: The Population Ahead, ed. Roy G.
Francis, pp.87-97. Minneapolis: University of Minnesota Press.
Experimental control of tree growth and reproduction. In: The
Physiology of Forest Trees, ed. K. Thimann, pp.601-10. New York:
Ronald Press.
Forsythia "Beatrix Ferrand." Natl. Hortic. Mag., 37: 112, 113.
Breeding ornamental trees and shrubs. Proc. Plant Propagators
Soc., 8:120-26.
1959
The cytogenetics of facultative apomixis in Malus species. J. Arnold
Arbor., 40:289-97.
1960
Meiosis in interspecific pine hybrids. For. Sci., 6:135-38.
Standing Room Only: The World's Exploding Population, rev. ed. (pa-
per). Boston: Beacon Press.
1961
With Hally Olivetti Sax. The effect of age of seed on the frequency
of spontaneous and gamma ray induced chromosome aberra-
tions. Radiat. Bot., 1:80-83.
Radiation sensitivity of Tradescantia microspore chromosomes to a
second exposure of X-rays. Radiat. Res., 14:66 '-73.
1962
With Hally Olivetti Sax. The effect of X-rays on the aging of seeds.
Nature, 194:459, 460.
Aspects of aging in plants. Annul Rev. Plant Physiol., 13:489-506.
1963
The stimulation of plant growth by ionizing radiation. Radiat. Bot.
3: 179-86.
With Lloyd A. Schairer. The effect of chronic gamma irradiation
on apical dominance of trees. Radiat. Bot., 3:283-85.
With H ally Jolivette Sax. The effect of chronological.and physio-
logical aging of onion seeds on the frequency of spontaneous
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KARL SAX
397
and X-ray induced chromosome aberrations. Radiat. Bot.,
4:37-41.
1964
The world's exploding population. Perspect. Biol. Med., 7:321-30.
Population problems. Topic, 8:5 -19.
1965
With H. I. Teas and Hally {olivette Sax. Cycasin: Radiomimetic
effects. Science, 149:541, 542.
1966
Biological problems of the age of science. Wash. State Rev., 10:5-
9.
The Bussey Institution: Harvard University's Graduate School of
Applied Biology. J. Hered., 57: 175-78.
With Hally Jolivette Sax. Radiomimetic beverages, drugs and mu-
tagens. Proc. Natl. Acad. Sci. USA, 55:1431 - 35.
Radiomimetic effects of beverages, drugs and insecticides. Cran-
brook Inst. Sci. Newsl., 36:46-49.
The world population explosion. Medicine Today, 1:8-14.
1968
With Hally Jolivette Sax. Possible mutagenic hazards of some food
additives, beverages and insecticides. {pn. I. Genet., 43:89-94.
With Hally Jolivette Sax and Wayne Binns. Radiomimetic effects of
veratrum. Toxicon, 6:69-70.
1969
Ethical aspects of the population crisis. BioScience, 19:303.
1970
With Hally Jolivette Sax and W. B. Itturian. Effects of sonic energy
on chromosomes. Environ. Mut. Soc. Newsl., 5:24, 25.
Representative terms from entire chapter:
karl sax
