Documenti di Didattica
Documenti di Professioni
Documenti di Cultura
Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at http://www.jstor.org/page/
info/about/policies/terms.jsp
JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content
in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship.
For more information about JSTOR, please contact support@jstor.org.
American Society of Plant Biologists (ASPB) is collaborating with JSTOR to digitize, preserve and extend access to Plant
Physiology.
http://www.jstor.org
This content downloaded from 128.187.103.98 on Mon, 02 Nov 2015 02:07:18 UTC
All use subject to JSTOR Terms and Conditions
DORMANCYAND GERMINATIONOF FBAXINUS SEEDS
George P. Steinbauer
This content downloaded from 128.187.103.98 on Mon, 02 Nov 2015 02:07:18 UTC
All use subject to JSTOR Terms and Conditions
814 PLANT PHYSIOLOGY
Observations
Structure of the seed and fruit of Fjzaxinus
Since the course of the germinationprocess depends largely upon the
structureof the seed, a brief descriptionof it is essential. In Fraxinus the
unit commonlyspoken of as the "seed" is in reality a winged fruit or
samara. The true seed is exposedonly after the ovary wall or pericarphas
been removed. The fruits, seeds, and embryosof red and black ash are
shownin figure1.
A B
Fig. 1. The fruit, seed, and embryoof Fraxinus. A. Red ash. B. Black ash. From
left to right: fruit, seed, embryo. Approximatelynormalsize.
Fig. 2. Crosssection of the fruit of black ash (diagrammatic). At the right a few
cells are shown from the layers of the seed external to the embryo. Cell contents have
been omitted.
P- pericarp EN- endosperm
SO- see(l coats M- mucilaginouslayer
SM- suberizedmembrane E- embryo
The seedcoats,SC, are madeup of five to eight layers of cells. The inner-
most cells of the seed coatsare separatedfrom the endospermcells, EN, by a
membrane,SM. The exact chemicalcompositionof the latter has not been
determinedalthough staining properties,solubility, and other microchemi-
This content downloaded from 128.187.103.98 on Mon, 02 Nov 2015 02:07:18 UTC
All use subject to JSTOR Terms and Conditions
STEINBAUER: DORMANCY AND GERMINATION OF ASH SEEDS 815
cal tests indicate the presenceof some cutin and suberin. This membrane
is one of the most resistant layers external to the embryo,and very likely
plays an importantpart in preventingrapid expansionof the embryo.
The reservefood materialsof the endospermconsist largely of fats and
reserveproteins. Starchand solublecarbohydratesappearin the endosperm
and embryoin the later stagesof the germinationprocess.
The embryo,E, is surroundedby a layer of cells, M, containingmucilagin-
ous material,calledthe ''schleimschicht'7by Lakon (9) relativeto the seeds
of Europeanash. Thesecells have thinnerwalls and less reservefood mate-
rial than those of the endosperm. Owingto the semi-gelatinousconsistency
of this layer, the embryosof Fraxinus seeds may be easily excised without
injury.
The degree of embryodevelopmentat maturity of the seed varies con-
siderablyamongthe differentspecies. The embryosare fully differentiated
into hypocotyl,cotyledons,and epicotyl,but the relative size of embryoand
seedvarieswidely. In figure1A thereis illustratedthe type of seed in w7hich
the embryoextendsthe full length of the seed. The seedsof red, white, and
green ash are of this type. Such seeds are quite easily germinatedif they
are first stratifiedfor a periodof one or two monthsat a temperaturenear 5°
C. In figure IB there is illustrated the type of seed in which the embryo
extendsfrom one-halfto two-thirdsthe length of the seed. Seedsof European
ash are said to be similar to those of black ash in this respect (9). Such
seedsdo not germinatereadily. They do not respondfavorablyto ordinary
stratificationtreatmentsand in nature often remainin a dormantcondition
in the soil for one or moreyears beforethey germinate(5, 9).
Dormancyand growth of excised embryosof black ash
Since the embryosof black ash are much smaller in proportionto the
rest of the seed than those of the other speciesstudied,it seemeddesirableto
makeobservationson themfirst.
In orderto determinewhetheror not embryosof recentlyharvestedseeds
are dormant,a numberof embryoswere excised from soakedseeds. They
were placed on moist blotting paper in Petri dishes and kept at room tem-
perature. At the end of two weeksthere was no evidenceof growthin such
excisedembryos.
Varioustreatmentswere then used in an attemptto overcomethis initial
dormancy. These treatments included different rates of aeration, use of
light, various substrata,and a series of differenttemperatures. The use of
higher than roomtemperatureswas effectivein inducing growth. Thus at
the end of four days someof the embryosin the 30°-C. chamberwere begin-
ning to show geotropiccurvatureof the hypocotyl. This was followed by
elongationof the cotyledonsand the formationof root hairs. "Whenthese
This content downloaded from 128.187.103.98 on Mon, 02 Nov 2015 02:07:18 UTC
All use subject to JSTOR Terms and Conditions
816 PLANT PHYSIOLOGY
days
1 0 0 0 0
2 0 0 0 0
3 0 0 0 0
4 0 0 0 6
5 0 0 0 6
6 0 0 0 8
7 0 0 0 10
8 0 0 0 16
9 0 0 0 22
10 0 0 0 24
15 0 0 8 44
20 0 0 12 49
25 0 0 18 60
30 0 4 18 60
This content downloaded from 128.187.103.98 on Mon, 02 Nov 2015 02:07:18 UTC
All use subject to JSTOR Terms and Conditions
STEINBAUER: DORMANCY AND GERMINATION OF ASH SEEDS 817
This content downloaded from 128.187.103.98 on Mon, 02 Nov 2015 02:07:18 UTC
All use subject to JSTOR Terms and Conditions
818 PLANT PHYSIOLOGY
chamber. Seeds were placed between layers of moist linen toweling and
stored at temperaturesof 5°, 20°, 25°, and 30° C. Embryoswere excised
from these differentlots of seed at the end of four, ten, twenty-eight,and
thirty-sixweeksand measuredfor increasesin length. The results obtained
are showngraphicallyin figure4. Measurementsmade on embryosheld at
25° C. wereintermediatebetweenthoseof the 20° and the 30° C. groupsand
havebeenomittedfrom the graph.
Thegrowthcurveduringthis periodof embryodevelopmentexhibitsthree
rather distinct phases. The slow growth at first may be accountedfor by
the initial embryodormancymentionedabove. This is followedby a period
of rapid elongationof both cotyledonsand hypocotyl. When the embryos
have attaineda length of 14r-14.5mm.,the rate of growthrapidly decreases.
Maximumsize is attained at betweentwo and three months at 20° C, and
considerablylater at 25° and 30° C. Further storage of seeds containing
enlarged embryosat temperaturesof 20° C. or higher does not ordinarily
result in germination. One lot of seed stored at 20° C. had fully enlarged
embryos at the end of two months and yet failed to produce a single
seedlingduring the followingtwo years of storageat 20° C.
That germinationof seeds with fully enlarged embryosis not due to
embryo dormancy is shown by the behavior of excised embryos. The
latter will often showmarkedgeotropiccurvaturewithin twelve hours after
excision,which is soon followedby elongationof cotyledonsand the forma-
tion of roothairs. By makinga longitudinalincisionin the seed,being care-
ful not to injure the embryo, one can often induce germinationwithin a
few days. This suggeststhat the delay in germinationis due to restrictions
placed upon the embryo by the enveloping layers: endosperm,suberized
layer, and seed coats. The pericarp probablyis not an important factor
since it disintegratesreadily at temperaturesof 20° C. or higher.
The retardinginfluenceof the envelopingtissueson furthergrowthof the
embryoseems to be primarily in the nature of mechanicalresistance. In
this connectionit is interestingto note that further growthof the cotyledons
is not retardedas much at the higher temperaturesas is the growth of the
hypocotyl. In this respectthe embryosare somewhatsimilarto the embryos
of Crataegus(3) and Ambrosia(2). Abnormalgerminationis not uncom-
monin the seedsof blackash and in otherspeciesof ash. The appearancesof
Fraxinus seeds exhibiting normal and abnormalgerminationare shown in
figure5.
It is interestingto note that 5° C. is not conduciveto embryoenlargement
althoughit is a very effectivetemperaturein forcing germinationafter the
embryoshave becomefully enlarged. Likewiseit is interestingto note that
whereas30° C. is a highly effectivetemperaturefor the growth of excised
embryosit is less effectivethan 20° C. when the embryosare still within the
seed.
This content downloaded from 128.187.103.98 on Mon, 02 Nov 2015 02:07:18 UTC
All use subject to JSTOR Terms and Conditions
STEINBAUER: DORMANCY AND GERMINATION OF ASH SEEDS 819
This content downloaded from 128.187.103.98 on Mon, 02 Nov 2015 02:07:18 UTC
All use subject to JSTOR Terms and Conditions
820 PLANT PHYSIOLOGY
J
I
i
III
GERMINATION
Most effective temperatures:Daily alternation of 20°-30° C.
Growthof excised embryosrapid at this temperature.
Supply of food accumulatedin embryosin stage II combinedwith high
temperaturesmakes possible sufficientpressure (osmotic, imbibitional,
etc.) to overcomemechanicalresistanceof layers external to embryos.
This content downloaded from 128.187.103.98 on Mon, 02 Nov 2015 02:07:18 UTC
All use subject to JSTOR Terms and Conditions
AND GERMINATION
STEINBAUER:DORMANCY OF ASH SEEDS 821
This content downloaded from 128.187.103.98 on Mon, 02 Nov 2015 02:07:18 UTC
All use subject to JSTOR Terms and Conditions
822 PLANT PHYSIOLOGY
This content downloaded from 128.187.103.98 on Mon, 02 Nov 2015 02:07:18 UTC
All use subject to JSTOR Terms and Conditions
STEINBAUER: DORMANCY AND GERMINATION OF ASH SEEDS 823
TABLE II
Vitality of Fraxinus seeds in relation to storage conditions
% % % %
Over concentrated sulphuric "]
acid 1.72 38 1.68 53
25 per cent, relative humidity 1 25° C. 5.55 46 5.36 48
50 " " " << 7.33 39 6.49 50
75 " " " li 11.39 0 9.85 1
5° C 7.50 40 7.30 54
20° C " 50 " 49
25° C " 43 " 56
30° C " 38 " 51
*
Storage of seeds for one year in sealed containers previous to planting.
t Figures based on quadruplicate samples of 100 seeds each.
Summary
1. A study of the optimumconditionsfor germinationof the seedsof red,
green, white, and black ash indicates that the seeds of black ash do not
germinateat temperaturesfavorableto germinationof the other species.
2. The differencesin temperaturerequirementsof the various species of
ash dependin part upon the size of embryoat maturity of the seed.
3. Embryosof black ash seeds undergoconsiderableenlargementbefore
germinationoccurs. Temperaturesnear 20° C. are most effectivein promot-
ing growthduring this period.
4. Germinationof the seeds of black ash does not occur at temperatures
favorableto enlargementof the embryo. Further growth of the embryois
retardedby mechanicalresistanceof the envelopingtissues and coats.
5. Stratificationof blackash seedsat 5° C. enablesthe fully enlargedem-
bryosto absorband accumulatereservematerialsfrom the endosperm. After
a periodof two or three monthsstratification,the embryosare able to over-
come the resistance of the enveloping membraneswhen placed at higher
temperatures.
6. Embryosof red, green,and white ash are fully enlargedat maturityof
the seed. Stratificationof the seeds at 5° C. brings about changesfavorable
to germination.
7. Dwarf seedlingsmay be obtainedfrom embryosexcised from freshly
harvestedseedsof all of the speciesstudied.
8. Storage of seeds in sealed containersat different temperaturesand
relative humiditiesshowedthat vitality of the seeds is retained the longest
This content downloaded from 128.187.103.98 on Mon, 02 Nov 2015 02:07:18 UTC
All use subject to JSTOR Terms and Conditions
824 PLANT PHYSIOLOGY
whenthe seedsare kept at a moisturecontentof less than 7.5 per cent, of the
dry weight. Seedswith a moisturecontentof less than 7.5 per cent, retained
their vitality equallywell over a wide range of temperatures.
9. Suggestionsare given for effectiveplanting proceduresfor the differ-
ent species studied.
University of Maine
Orono, Maine
LITBEATUEE CITED
1. Crocker, Wm. Mechanicsof dormancyin seeds. Amer. Jour. Bot.
3 : 99-120. 1916.
2. Davis, W. E. Primary dormancy,after-ripening,and the development
of secondary dormancyin embryos of Ambrosia trifida. Amer.
Jour. Bot. 7 : 57-76. 1930.
3. , and Rose, R. C. The effect of external conditionsupon
the after-ripeningof the seeds of Crataegusmollis. Bot. Gaz. 54 :
49-62. 1912.
4. Findeis, M. Tiberdas Wachstumdes Embryosim ausgesatenSamenvor
der Keimung. Sitzungsber.kaisl. Akad. Wiss. Wien 126 : 77-102.
1917.
5. Heritage, W. Black ash. Jour. For. 34 : 531-533. 1936.
6. Jones, H. A. Physiologicalstudy of maple seeds. Bot. Gaz. 69 : 127-
152. 1920.
7. Lakes States For. Exp. Sta, Recent results from seed dormancytests.
For. Res. Digest 1-2. May, 1935.
8. . geed treatmentfor shelterbeltspecies. For. Res.Digest
6-7. Aug., 1935.
9. Lakon, G. Beitragezur forstlichenSamenkunde. II. Zur Anatomie
und Keimungsphysiologieder Eschensamen. Naturwiss.Zeitschr.
Forst- und Landw.9 : 285-298. 1911.
10. Puchner, H. Die verzogerteKeimungvon Baumsamereien.Forstwiss.
Centralbl.44 : 445-455. 1922.
11. Yerkes, G. E. Propagation of trees and shrubs. U. S. Dept. Agr.
Farmers' Bull. 1567. 1929.
This content downloaded from 128.187.103.98 on Mon, 02 Nov 2015 02:07:18 UTC
All use subject to JSTOR Terms and Conditions