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VOLUME LIX NUMBER 6

THE

BOTAN ICAL GAZETTE

JUNE -1915

A STUDY OF DELAYED GERMINATION IN ECONOMIC

SEEDS
CONTRIBUTIONS FROM THE HULL BOTANICAL LABORATORY 204

DEAN H. ROSE

(WITH ONE FIGURE)

This paper presentsthe resultsof an attempt to discover some

of the practicalproblemsthat seedsmenand growershave to meet,
and to work out, so far as possible, practical methods of solving
these problems. The seeds tested were furnishedby six of the
leading seed houses of the United States.
In the presentstate of our knowledgeit can be said that delayed
germinationand poor germinationare due to one or more of the
following causes: hard-coatedness, the need of after-ripening,
exclusion of oxygen by the seed coat, the effectof froston seeds,
fungion or in seeds, and of course the presenceof seeds containing
dead embryos.
Hard-coatedness
The condition of hard-coatedness in the seeds of legumes is
well known. To overcome this conditioninvestigatorshave used
hot water, chemicals, and mechanical devices for scratching or
puncturingthe seed coat.
The use of hot water for forcinggerminationis undoubtedly
older than the referencesto it in periodical literature. It was
recommendedby BRUYNING (6) in i893 forseeds of Ulex europaeus,
425
426 BOTANICAL GAZETTE [JUNE

and by WERNICKE (30) in i895 forseveral different kinds of seeds.

Mention may also be made of the work of JARZYMOWSKI (I7) in
I905 with seeds of various economic legumes, and of BOLLEY (3)
in I9I2 with those of alfalfa. BOLLEY obtained positive improve-
ment in germinationif exposure to a given temperaturewas not
long enough to kill the embryo.
Treatment with concentrated sulphuric acid dates from the
workof ROSTRUP(25) in i896-i897. It was also used by TODARO
(29) in i9oi, by HILTNER (14) in I902, by JARZYMOWSKI (I7) in
I905, and by BOLLEY (3), and LOVE and LEIGHTY (22) eight years
later. Increased germinationwas obtained in all these cases.
Treatment with other chemicals has included the use of ether,
ethyl, and other alcohols (VERSCHAFFELT3I, I9I2), chloroform,
sodium hydroxide,postassium hydroxide,potassium nitrate, and
mercuricchloride. Of these, the lower alcohols are the only ones
that are very effective.
There are obvious practical objections, however,to the use of
eitherhot water or chemicals. As a consequence, therehave been
numerouseffortsto devise means forthe mechanical treatmentof
hard-coatedseeds. In Germany,KUNTZEand Huss (i6), working
about i890, were able with a scratchingmachine to increase the
germinationof Lathyrussylvestris 83 per cent, Vicia Cracca 7I per
cent, and AstragallusGlycyphyllos 77 per cent.
Somewhat later MICHALOWSKIdevised an apparatus in which
the seeds were passed between two rollers,one of rubber,the other
of rough steel. Smaller sorts of seeds were badly crushedby such
a mechanism,and it was later displaced by two others,one designed
by the WissingerSeed Co., of Berlin, the other,called a "prepara-
tor," by H. NILSSON of the experimentstation at Svaldf, Sweden.
In both of these the seeds are thrownfroma revolvingdisk against
the concave surface of a circular rough stone, within which the
disk revolves. HUME and GARVER (I5), using the "preparator,"
obtained a definiteincreasein the germinationof seeds of Medicago
sativa,M. media, and MI.falcata. Anothermachine now in use in
England has made it possible, according to CARRUTHERS (7), the
designer,to buy clover seed guaranteed to germinate98-ioo per
cent. The seeds to be treated are fed into a revolving cylinder
I9I5] ROSE-DELA YED GERMINATION 427

lined with sharp, close-setsteel points against which the seeds are
thrownand scratchedas the cylinderrevolves.
Mention should be made here also of an apparatus invented
by KtHLE (ig) for scraping the rough outer covering from sugar
beet "seed." Very satisfactoryresultshave been obtained by its
use, since "seeds" so treated absorb water better than untreated
ones, and germinatemore rapidly; they also give a better total
germination,on account of the removal of fungus-infected mate-
rial fromthe outside of the "seed," especially if this removal is
followedby treatmentwith some fungicide.
With any one of the machines here described except the last,
whichserves a slightlydifferent purpose,it has been founddifficult
to treat every seed that passes throughand, at the same time, to
avoid serious cracking of the coat or bruisingof the entire seed
(GLOCKENTOEGER I I).
It is believed that these difficultieshave been avoided in a
machine devised and in use duringthe winterof I9I2-I9I3 at the
Hull Botanical Laboratory of the University of Chicago. This
machine consists of a direct pressure blower, furnishedby the
ConnersvilleBlower Co., to whichis attached an apparatus through
which seeds can be fed and blown against the points of a bank of
needles. In experimentsconducted with this machine,the blower
was drivenby a two horse-powermotorand gave pressuresas high
as 2 .5 pounds to the square inch. The needles used were of three
sizes, nos. 4 and i i sewingneedles and no. 4 darningneedles,made
up into three different cylindricalbunches or banks, each bank of
course consistingof only one size of needles. The needles were
held togetherby solder at the eye end and by wireor a ferruleone-
half to two-thirdsof the distance fromthe eye to the point.
In the cut here shown (fig. i) the needles are about half an
inch fromthe end of the air tube. In practice a screen cap is
placed over the needles and the tube as a coveringfor a glass jar
beneath, into which the seeds fall. To use the apparatus, valve
e is closed and valve b is opened; seeds are poured into compart-
ment c; valve b is closed and the blower started; valve e is then
opened wide enough to let the seeds out, but not so wide that they
interferewith each other as they strike the needle points. It is
428 BOTANICAL GAZETTE [JUNE

plain that the distancethe valve is to be openedwill vary with

differentseeds,but will not be at any timeparticularly hard to
determine.Withvalve b closed,therewas no difficulty in getting
the seeds down into the air tube; withit open theywould be
blownout at d.
The openingat h is five-eighthsofan inchin diameterand will
accommodateseedsof sweetpeas, lupines,Lathyrus, honeylocust,

C,

e~~~~~~~~~~~~

ing themagainst a bank of needle points.

etc. When smaller seeds are to be treated, an attachment with

inchfeed,valves,and airtubecan be adjustedreadily.
three-eighths
Valveg can be usedto regulatethepressure,
as readon thepressure
gaugef
Testsweremadeof treatedand untreatedseeds,on filterpaper
kept moist withdistilledwater,at a temperatureof23.25' C. The
resultsare summarizedin table I.
I915] ROSE-DELA IYED GERMINATION 429

It is plain that treatmentwith the machine increased germi-

nation considerablyin the case not only of legumes, but also of
snapdragon, Delphinium, sweet marjoram, Ipomoca, okra, and
lettuce.
TABLE I
GERMINATION OF SEED TREATED WITH MACHINE

GERMINATION
KIND OFOF
SEED DURATION OF __ _ __
KINO SEED
~~TEST
IN DAYS -
Untreated Treated

Alfalfa............................... IO 5I 99
Alfalfa............................... I0 74 94
Bossiaea heterophylla .................. 30 3 73
Bossiaea scolopendria.................. 30 II 52
ClianthusDampieri.................... IO 0 80
White clover. .......................... 10 70 94
Delphiniumchinense................... 30 74 96
Dilliwyniaericifolia .................... 30 2 77
Gleditschia........................... go 0 70
Ipomoea (average of 3 sPP.)............ IO 34 71
Lathyrus(average of 3 vars.). I4 63 93
Lettuce (average of I I vars.) ...... ...... I2 5 83
Sweet marjoram....................... 30 54 96
Mustard (average of 2 vars.).14 20 69
Okra ................................ 30 34 79
Perennialpeas (average of 2 vars.) ....... Io 62 87
Sweet peas (average of 5 vars.) .......... 14 70 99
Platylobiumtrilobatum. 30 I3 64
Snapdragon ........................... 30 6 1 23
Vetch................................ IO 77 92
Average . . 40.7 83.7
Increase due to treatment ... ............ .I . 43 7

It is not strictlycorrect,however, to call lettuce seeds hard-

coated. That theirgerminationis improvedby treatmentwiththe
machine shows delay to be due to coat restrictions. The coat
restrictionsare removed also by soaking in water for 24 hours,
hence it is likely that delay is caused by a reduction in rate of
water absorption rather than by lack of oxygen. Whatever be
the character of the seed coat, or coats, which interfereswith
germination,it disappears as the seed grows older. These points
are well illustratedby table II. The data here presentedsupport
the statementmade by various seedsmen,that two or three-year-
old lettuce seed gives better germinationthan fresh. How this
condition comes about is not known, but it probably depends on
430 BOTANICAL GAZETTE [JUNE

changes in the permeabilityof the seed coat. It is certainlynot

a matter of embryo changes.

TABLE II
GERMINATION OF TREATED AND UNTREATED LETTUCE SEED

GERMINATION

V'ARIETY DURATION OF
TEST IN DAYS Treated with Soaked in H.O
Untreated machine 24 hrs.

Black-seededbutter-
I909 ................... I 2 98 ............ ............
I910 ................... I2 99 ............ ............
19I .1. . . . . . . . . . . .... I2 89 93 95
I912 .1................. 12 67 97 100
Prize head-
I909 ................... 12 98 ............ .
I910 ................... 12 97 ............ ............
I9I I. .................. 12 84 96 97
I912 ....... ............ 12 76 98 96

It appears, froma study of the recordskept in this work,that

not only the total germination,but also the rapidityor energyof
germinationis greaterin treated than in untreatedseeds. This is
shown in table III.
TABLE III
RAPIDITY OF GERMINATION OF TREATED AND UNTREATED SEEDS*

Kind of seed Germination after Untreated Treated

Alfalfa...................... 3 days 48 98
White clover.................. 3 * 69 89
Perennialpeas . . 4 0 33
Perennialpeas ... . 4 24 8o
Lupines..................... 4 40 88
Delphinium.................. 5 4 44
Sweet peas . 4 7
Sweet peas .................. 4 i6 86

* For final germination percentage of these samples see table I.

Such rapidityof germinationwould clearlybe of advantage in

making the crop uniformin size and age, and in keeping down
weeds. Indeed, it may be said truly that vigor of germination
and vigor of the plants produced are more importantthan merely
high germinationpercentage. Plants that get a good strongstart
1915] ROSE-DELA YED GERMINATION 43 I

are more certain to be productive than those that for any reason
are weak fromthe beginning.
Before this machine can become commercially practicable,
experimentsmust be conducted to determine: (i) the possibility
of substitutingsomethingelse for needle points; (2) the proper
distance these points should be fromeach other to give the best
resultsfordifferent sized seeds; (3) the pressurenecessaryto give
the best germination for differentkinds of seeds; for certain
legumes a pressureof two to threepounds is necessary,forlettuce
one pound or even less; (4) the effectof storageon the germination
of treated seeds; (5) the germinationof treated seeds in soil. To
be effectivein overcominghard-coatedness,the needle point need
only pass throughthe palisade layer and not entirelythroughthe
coat. Even withthisslightdeformationit is possible that bacteria
and fungi can gain an entrance. That destructionby bacteria
and fungi actually does take place was shown by JARZYMOWSKI(I7)
forseeds of Ulex europaea, lupines, and other large-seededlegumes
which had been treated with the Wissingermachine. Red clover
and Lotus corniculatuswere the only ones whose germinationin
soil aftertreatmentwas not seriouslyreduced.
As to the germinationin soil of seeds treated by the blowing
method here described, there are not at present enough data on
hand to justifythe drawing of definiteconclusions. Preliminary
experimentsseem to indicate for alfalfa seed, where the percent-
age of hard seed is high,that germinationin soil is definitelybetter
after treatmentthan before. Further investigation,of course, is
necessarybeforethis can be confirmed. In conclusion it may be
stated that there was no serious crushingor crackingof seeds or
seed coats by this machine.

The need of after-ripening

This is a conditionwhich occurs, to mentiononly a few cases,
in seeds of Crataegus,various conifers,Fraxinus, potato tubers,
and lily-of-the-valleybulbs. For a discussion of the general situ-
ation and a resumeof the literaturethe reader is referredto the
paper by ECKERSON (io) dealing with after-ripening in the seeds
of Crataegus. The work to be discussed here had to do with the
432 BOTANICAL GAZETTE [JUNE

germinationof seeds of conifers,and specificreferencewill be made

to a fewof the more pertinentpapers on the subject.
It is a matter of common knowledgethat coniferseeds germi-
nate slowly. It is also well known for several of them that as
they grow older the rapidity of germinationincreases, up to at
least the end of the first 6 months after they were gathered.
SCHWAPPACH(27) states that in the fallseeds ofA biesdid not begin
to show sprouts for 6o days, and required4o days more beforethe
test could be considered closed. In March they began almost
immediatelyand finishedin 20 days. The conclusion is natural
that after-ripeningtakes place, and this, in fact, is assumed by
workerswho have recentlyattacked the problem. HILTNER and
KINZEL (14), it is true,reasoningfromresultsobtained by treating
seeds of Pinus Strobus,P. Peuce, and P. Cembrawith concentrated
sulphuric acid, ascribed the delay to coat restrictions. LAKON
(20) has made the objection that the tests on which these authors
relywere too fewand on too small a numberof seeds. He repeated
their experimentswith the same three species of pine, but could
obtain no forcingof germination. Untreated seeds took up water
just as well as did the treated ones, even though theirouter coats
were hard. Careful determinations of the amount of water
absorbed by untreated seeds of Pinus sylvestris,P. Strobus, P.
Peuce, and P. Cembrashowed that all of them reached nearly the
maximum in 24-48 hours. Increases in weight after that time
were practicallynegligible; hence it is clear that such seeds cannot
be considered "hard-coated" like the seeds of legumes. More-
over, the cutting test, applied to these seeds, showed that all of
themweredamp, that is, had absorbed water. Increases in weight,
therefore, were not due to a feweasily swellingseeds. From these
resultsLAKON concludes that coniferseeds are not, strictlyspeak-
ing, "hard-coated," and that delay is due to conditions within
the embryo.
AlthoughLAKON found concentratedsulphuricacid ineffective,
CORREVON states, in a paper published somewhat earlier, that
weak acid (o. 25 per cent acetic or 2 per cent phosphoric)increases
the germinationof seeds of JuniperusCedrus.
SCHWAPPACHrecommendscold storage for 14-30 days (he does
not say how cold) forseeds of Pinus Strobus,followedby a germi-
1915] ROSE-DELA YED GERMINATION 433

nation temperatureof 25 C. The common practice of layering

various coniferseeds doubtlessfindsits justificationin a shortening
of the time requiredforgermination. Low temperatureshave also
been used forpreservingthe vitalityof coniferseeds. HAACK(I2)
driedseeds so theylost about 2 per cent in weight,then storedthem
in dry, air-tightcontainers on ice. CLEMENS (9) stored seeds in
the refrigeratorof a brewery in vessels containing sodium car-
bonate to absorb moistureand carbonic acid. Both investigators
report that seeds thus stored remained viable longer than those
kept under ordinarylaboratory conditions.
In order to analyze the situation more carefully,the following
series of experimentswere conducted on seeds of various conifers.
i. Tests of untreatedseeds.

2. Tests of seeds which had been in cold storage (3-5' C.):

(a) in wet sand, (b) in weak solutions of hydrochloricacid, (c) in
distilled water.
3. Tests were made with seeds which had been injected with
weak hydrochloricacid or with water, by exhaustingthe air from
them when they were in these liquids, and then restoringthe
pressureto normal. This was repeated at least threetimes forall
seeds here spoken of as injected. Table IV summarizesthe results
obtainedin series i and 2a.
TABLE IV
GERMINATION OF CONIFER SEEDS, UNTREATED AND AFTER STORAGE IN WET SAND
AT 3-,5 C.; PERCENTAGE GERMINATING AFTER ONE MONTH IN GERMINATOR

DRY STORAGE UNTREATED COLD WET STORAGE FOR

2 3 4
month months months months
KIND OF SEED | _ - -

Date of starting germination

Jan. 26 April 4 April 24 May 30 Feb. 26 Mar. 25sApril 26 May 28

Cupressusmacrocarpa... 8 a I ...... I5 27 40 36
Picea Menziesii. . ..............56
42 ...... 23 70 6o
Pinus austriaca.... 57 48 27 28 88 64
Pinus Strobus.......... 12 10 6 28 34 40 44 59

The results given in the table show that germination was

definitelyincreasedby cold wet storage for fourkinds of conifers.
434 BOTANICAL GAZETTE [JUNE

In no case does the maximumgerminationof seeds of a givenkind

from dry storage equal that of seeds of the same kind fromcold
wet storage. The effectof the cold wet storageis most noticeable
in the cases of Pinus Strobus and Cupressus macrocarpa,where
increasesof 32 per cent and 3I per cent respectivelywere obtained.
For all of the seeds here reportedon, except those ofPinus Strobus,
dry storage seemed to cause a decrease in viability. This can be
seen from the first four columns of the table. Tests of Pinus
Strobus were run for 6o days, the other three for 3o days.
TABLE V
GERMINATION OF SEEDS OF CONIFERS AFTER SOAKING IN WEAK ACID AND
STORAGE AT 3-5 C.

ACID (HC1) WATER

Soaked Injected
KIND OF SEED AND LENGTH OF CON- |
TIME IN COLD STORAGE TROL I
o 0 0 0 0 Soaked Injected
o 0
0
0

Pinus Strobus-
3 days.44 58 15 66 54 .
6 .. 5' 57 4
IO ................. .... 53 65 67 1 63 ......
N o cold storage .......... 30 ...... ...... ...... ...... ...... 68
Pinus austriaca-
3 days . ....... ...... 64 .... 52 ...... .. 65 ..
6 . ........ ...... 48 42 ...... I ......
59 ......
10 ......... 35 L... 50 ... ...... 45 ......
No cold storage......... 52 54 ...... ...... .. ......

The series of experimentsshown in Table V was planned to

determinewhetherdelay in germinationis due to an alkaline or
neutralreactionof the embryo. It was thoughtthat if such is the
case, weak acid solutions would change the reaction sufficiently
to cause growthto begin,when the seeds were placed in the proper
conditions.
The results obtained for Pinus Strobusdo not, however,bear
out this theory. Seeds injected with distilled water gave i8 per
cent bettergerminationthan those merelysoaked in it, and slightly
betterthan those injected with weak hydrochloricacid. It seems
likely fromthis that delay is due merelyto lack of water. When
1915] ROSE-DELA YED GERMINATION 435

this water was supplied, by long soaking or by forcibleinjection

under pressure,germinationwas much improved.
Results for Pinus austriaca are less conclusive on this point.
They do show, however,and the same is true forP. Strobus,that
soaking in eitherwater or weak acid gave greatergerminationthan
was obtained in the controls,38 per cent and 13 per cent respec-
tively. For P. austriaca,better results were obtained fromshort
than fromlong soaking.
Referringagain to table IV, it is possible that the increases in
germinationshown therewere due not so much to the cold storage
in itself as to the thoroughinfiltrationof the seeds with water.
There is need of much more work on this question before any
definiteconclusionscan be drawn.

Exclusion of oxygen
No attempt will be made to review formerwork, since this
has already been done by SHULL (28). Results presentedin table
VI seem to indicate that the germinationof certain economic
seeds is delayed for lack of oxygen. They also indicate the need
of a detailed studyof these seeds.
TABLE VI
GERMINATION OF SEEDS TREATED WITH OXYGEN OR HYDROGEN PEROXIDE

Kind of seed
Kind of Duration of Untreated
Uteed In 8o per cent In 0I 5 per cent
seedtest in days oxygen M~o.

Dandelion 6................ 14 56 ... I........ 72

Datura-
Golden Queen .......... 14 73 ............ 70
W rightii............... 14 20 ............ I00
Lettuce-
Grand Rapids 0.......... I 44 32
Martynia ........ ........ 20 0 90 8o

Datura Wrightiiwas forcedconsiderablyby hydrogenperoxide,

Datura Golden Queen not at all. Lettuce gave good results,but,
as has been suggested,this is probably due to absorptionof water.

Effectof froston seeds

ATTERBERG (2) says that seeds of oats and barley harvested
in Sweden aftera heavy frostgave fairto good germinationin the
436 BOTANICAL GAZETTE [JUNE

laboratory,but in many cases no plants when sown in the field.

Unpublished data obtained by EASTHAMin Canada show that the
germinationof oats grownin the prairieprovincesis oftenseriously
reduced by early frosts. He says, "as far as our observationsgo,
a couple of degreesof frostin the milk stage are in many instances
sufficientto ruin oats for seed. In the dough stage they are not
nearly so susceptible, and when well ripened and dry stand con-
siderable frost without serious injury." EASTHAMfound also
that such seed, germinatingpoorlywhen harvested,oftenimproved
with age. This seems to indicate in such cases the necessityfor
a period of after-ripening. Through the courtesyof the Canadian
seed laboratoryand two Americanseed houses the author has had
the privilege of testing several samples of frosted oats. The
resultsare summarizedin table VII.
TABLE VII
GERMINATION OF OATS

HULLS REMOVED; ON COTTON IN PETRI DISHES

UNTREATED; UNTREATED; HULLS OFF; AT 22' C.
ON FILTER SENDER'S ON FILTER _ _ __
VARIETY
ARTY PAPER AT TEST PAPER AT
20?C. I8?-20' C. 20? C. 8o per cent 6o per cent 40 per cent 20 per cent
.._ - Ioxygen oxygen oxygen oxygen

Lincoln 85 40 79 ......... ...... ......... .........

Swedish. 77 40 8o ...... ......... .... .........
4920 .... 34 35 38 | 31 25 38 33
5139 .... 46 ......... ......... ......... ......... ......... .........
34 77 .... 56 43 .... ......... ......... ......... .........
3974.... 45 ! 42 ......... ......... ......... ......... .........
4948.... 89 62 94 1 93 97 95 96
5302 . .. . i6 i 27 .. ..... .. ... .. .. . . .. . .. .. ..... . .. . .I . . . .. .

The best germinationwas obtained fromhulled seeds in oxygen,

though the results are more clear-cut for no. 4948 than for no.
4920. No definiteconclusions can be drawn as to what percent-
age of oxygenis mosteffective. It is noteworthythat two samples,
nos. 3477 and 4948, show much better germination,23 and 27 per
cent respectively,than they did when tested in the Canadian Seed
Laboratory six months earlier. This agrees well with the state-
ment made above that frostedoats go througha process of after-
ripeningand improvein viabilityas theygrowolder. There is the
same need of after-ripeningin wild oats (Avenafatua), as has been
19I51 ROSE-DELA YED GERMINA TION 437

shown by ATWOOD. But referringto the table again, it will be

seen that one sample, no. 5302, deterioratedin vigor as it grew
older, since it gave a percentageof 27 when firsttested and i i per
cent less 6 monthsafterward. Consideringthe resultsas a whole,
it is clear that frostedoats are unreliable in performanceand of
very doubtfulvalue forseeding purposes.
Another crop which sometimes suffersfrom frost is garden
peas. Within the last two or three years the growingof garden
peas for seed has become an important industry in Idaho and
Montana. It has been found that certain of the late varieties
grown there are injured by frost and the viability of the seed
seriouslyimpaired.
A study of 14 samples of such peas has shown that decrease in
germinationis probably due to two different causes, both of which,
however,may be the effectof frost.
i. Actual injuryto the embryo,especially the tip of the radicle.

It has a whitish shriveled appearance and starts to grow very

slowly,if at all.
TABLE VIII
GERMINATION OF PEAS

Variety
Variety I ~~Coats
Coats200
on;. Coats on;
25
Coats off;
20'

PremiumGem........................ 90 82 86
Nott's Excelsior 73I86 .................
7
71531
I ................. 98 100 .
Telephone 885H....................... 58 46 1-
GradusQI3K ......................... 44 36 58
9 13S . . . ....................... 70 ............ 76
Dwarf Defiance 874C.................. 56 ............ 52
"t '. ............
"t 874K.64 96
"t 874H.................. 78 ............ 76
Alderman 912S ........................ 66 ............ ...........
912K ....................... 82............. 96
" 912T ....................... 56..... 5 6
912V .78........ 78 ............ 9)
Telegraph 68528 ....................... 48 ............ so
Average.........6.......... . 67 ........... 74

The presence of fungi on or in the seed coat. That this

2.
actually decreases germinationwas shown by the work summar-
ized in table VIII. With the coats on, the seeds of all the samples
here reportedon showed much fungus infection; with coats off,
438 BOTANICAL GAZETTE [JUNE

very little or even none at all. Ten samples gave an average

germinationof 67 per cent with the coats on, and 74 per cent with
the coats off,a differenceof 7 per cent. Individual samples, such
as Dwarf Defiance 874K, gave even more strikingresults.
The condition of garden peas, with reference to fungi, is
approached more or less closely by that of a large numberof other
garden and flowerseeds as is shown in table IX, summarizingthe
generalresultsof thisinvestigation,and in the discussionfollowing.
Plants whose seeds show delayed germination,classifiedaccord-
ing to probable causes (the word "probable" is used advisedly,
forwhile the evidence is convincingin some cases, it is much less
so in others):
i. Hard-coatedness.-Canna, ClianthusDampieri, Delphinium,

Erythrina,Hibiscus, Ipomoea (4 spp.), Lathryus,Lupinus, sweet

peas (4 vars.), snapdragon, alfalfa, sweet clover, white clover,
lettuce (io vars.), mustard (2 vars.), okra, sensitive plant, sweet
marjoram,vetch,Gleditschia.
2. Frosted.-Oats, peas (8 vars.).

3. Need of after-ripening.-Wild cucumber, Picea (3 spp.),

Pinus (2 Spp.).
4. Exclusion of oxygen by the seed coat.-Datura Wrightii,
Martynia.
5. Cause of delay not determined.-Coix Lachryma, feather
grass, Pampas grass, asparagus, barley,blue grass,cardoon,celery,
chives, dill, horehound,kaffircorn, leek, millet, parsley, parsnip,
pepper, radish, rosemary,spinach, summersavory, thyme,Aqui-
legia, Asparagus Sprengeri,Bignonia, Centaurea,Clematis,dande-
lion, Datura Golden Queen, Eschscholzia, foxglove, heliotrope,
Helianthus, hop, lavender, Momordica, Nasturtium, Oenothera,
pansy, Pentstemon,Primula, Salvia, Verbena,Abies Mertensiana,
A. pectinate,Berberis,Betula alba, Cupressushorizontalis,C. macro-
carpa, C. pyramidalis, Larix. Further work would doubtless
explain the cause of delay in many of these seeds and make the
growingof plants fromthem a much simplermatterthan it now is.
Verypertinentat this point are the resultsfromthe Minnesota
Seed Laboratory for i9io and i9ii (OSWALD 23). Of field seeds
I4 kinds were tested; of garden seeds 26 kinds. Of fieldseeds, for
19151 ROSE-DELA YED GERMINATION 439

00

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440 BOTANICAL GAZETTE [JUNE

3 of the kinds, So per cent or more of the samples were below the
governmentstandard of germination. Of garden seeds, for i6 of
the 26 kinds, 50 per cent or more of the samples were below stand-
ard. Mention should be made also of work by BROWN (5) on the
germination of packeted vegetable seeds. He found that the
average germinationof box vegetable seeds put up by 6o firmsfor
four years was 6o. 5 per cent. The lowest average for any firm
was 36. 5 per cent, the highest8i . 5 per cent. The average germi-
nation of packeted vegetable seeds put up by 20 mail-orderhouses
in i9ii was 77. 5 per cent (lowest average 76. 2 per cent, highest
77. S per cent). Justwhat these figuressignifyis not clear. There
are threepossibilities: (i) the seeds werepoor because of the seeds-
man's dishonestyor carelessness; (2) the seeds were poor because
it is not possible with present methods to produce better ones;
ifso, the governmentstandardis, now at least, too highand methods
of production need improvement; (3) the seeds seemed poor
because presentmethodsof makinggerminationtestsdo not always
adequately determinethe value of a given sample. In the writer's
opinion,the responsibilityforlow test mustbe sharedabout equally
by all three,thoughthe firstis a less importantfactorthan it was
a fewyears ago.
6. Plants whose seeds were found infected with fungi.-
Feather grass, asparagus, beggarweed, buckwheat,cardoon,celery,
chives, sweet clover, dill, kaffircorn, leek, millet, oats, parsley,
peas (I2 vars.), pepper, radish, rosemary,spinach, thyme,vetch,
Aquilegia, Asparagus Sprengeri, Bignonia, Clematis, Clianthus
Dampieri, wild cucumber, dandelion, Datura Golden Queen, D.
Wrightii,Helianthus, hop, Ipomoea (4 spp.), Lathyrus,lavender,
Nasturtium,pansy, sweet peas (4 vars.), Primula, Verbena,Abies
Mertensiana, A. pectinate, Berberis, Cupressus horizontalis,C.
macrocarpa,C. pyramidalis,Picea excelsa,P. Menziesii, P. rubra,
Pinus austriaca,P. Strobus.
The speciesand varietiestestedwere134, but 30 of theseare
omittedfromthe second sectionof table IX; 29 of thesegerminated
rapidly and well, and one other,on account of bad infectionwith
fungi,showed not delayed but definitelypoor germination; 69, or
5I .4 per cent of the total, were found more or less infectedwith
'9Is] ROSE-DELAYED GERMINATION 44I

fungi. This point was determinedforeach kind of seed not more

than two days after the test began. All filterpaper was boiled
5-IO minutes before being used and kept moist with distilled
water during the test. Repeated washing of seeds and removal
to freshfilterpaper showedthatin all cases infectioncame fromthe
seeds, not fromthe paper.
There is no intentionhere of implyingthat seedsmenin general
purposelyput on the marketseeds low in vitalityor badly infected
with fungi. It does seem clear, however, that there is need of
closer supervisionby the seedsmen themselvesof all stages of the
process of seed production; alternation of crops to avoid soil-
infection, cultivation, harvesting, threshing, cleaning, storage;
all of these need close attentionif seed of the best quality is to be
produced. The most candid way in which to approach the whole
question is to admit that seed analysts, seed-growers,and seed
merchants do not at present know a numberof thingsthey need
to know in referenceto the question of fungusinfectionof seeds,
and to all the other questions consideredin thispaper. The whole
matter constitutesan extremelycomplex physiologicaland patho-
logical problem,with verypractical aspects, the solution of which
can be broughtabout only by carefulstudy fromseveral different
points of view. To be specific,the followinglines forinvestigation
may be suggested:
i. The relationof germinatortests to the actual vegetation of

seeds in the soil. This should be studied througha periodof several

years.
2. The relationof fungion or inside of seeds to the germination
of such seeds in soil. At the riskof seemingto repeatunnecessarily,
the writerwishes to say that in his opinion the importanceof this
problem is only poorly appreciated in this country. Some recog-
nition of the dangers accompanying fungusinfectionof seed has
appeared of late in the work of BOLLEY and others in the United
States, and in German agriculturalliterature. APPEL (i), writing
on the relation of pathology to seed control, says that in seed-
testingstations,pains should be taken to give judgment as to the
presence of spores of plant diseases on seeds to be examined.
It is his opinion, further,that in comparative field tests more
442 BOTANICAL GAZETTE [JUNE

attentionmust be given than formerlyto pathologicalphenomena.

Observationson this point should be given along with other data
fromthe experiment.
3. The causes of delayed germinationin asters, certain hardy
perennials,labiates, ornamentalgrasses,cucurbits,conifers,frosted
oats, Betula, and Berberis.
4. The value of hard seeds of legumeswhen planted in the soil.
5. The relation of any or all of the causes of delayed germi-
nation to the vigor of the plants produced. It is not enough that
a given lot of seeds shall be free fromimpurities; it is not even
enough that it shall give a high germinationpercentage. It must,
above all, give rise to vigorousproductiveplants, when planted in
field conditions. Consequently, any knowledge which will teach
us how to grow such seeds and how to know poor seeds is of the
greatestpractical importance.

Summary
Hard-coated seeds of legumes, and seeds of Delphinium,
i.

Ipomoea, lettuce,mustard,okra, sweet marjoram,and snapdragon

can be forcedto more rapid germinationby being blown against
needle points.
2. For two varietiesof lettuceit is shownthat the seed improves

in viability as it grows older, up to the end of at least the fourth

year. This improvementis probably due to increased perme-
abilityof the innerseed coat to water.
3. Cold storage in wet sand increased the germinationseeds of
Pinus Strobusby 32 per cent, of Cupressus macrocarpaby 3I per
cent. Delayed germinationof coniferseeds, more especially those
of Pinus Strobusand P. austriaca,seems to be due to lack of water
intake, and not to an alkaline or neutral reaction of the embryo.
This statementis supported by the fact that seeds injected with
distilledwater gave better germinationthan those merelysoaked
in water or in weak acid at the temperatureof meltingice. Any
kind of soakingor injectiongave I3-38 per cent bettergermination
than was obtained with the controls.
4. Certain samples of frosted oats improve in germinating
power as theygrowolder,othersdeteriorate.
1915] ROSE-DELAYED GERMINATION 443

5. Certain late varieties of western-grown garden peas germi-

nate poorly. This is shownto be due to one or both of two causes:
(a) actual frostinjury to the embryo; (b) the presence of fungi
on or in the seed coat or inside of it.
6. Seeds of 5I .4 per cent of all species and varieties examined
showed fungion the seed coat withintwo days afterbeing put to
germinate.

The writer is indebted to Dr. WILLIAM CROCKER for many

suggestionsand criticisms.
UNIVERSITY OF CHICAGO

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APPEL, O.,
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Vers. Stat. 60:427-432. I904.
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5. BROWN, EDGAR, The germinationof picketed vegetable seeds. U.S.
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