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THE SPUR SHOOT OF THE PINES
ROBERT BOYD THOMSON
the leaves are numerous, while in Pinus they are much more
restrictedin number,not more than 8 having been recorded up
to the present. In the former,too, the numberis indefiniteand
the leaves are spirallyarranged,while in Pinus the definitecyclic
arrangementhas been established. A parallel to this is seen in the
angiospermflower,where the lower formsof both monocotyledons
and dicotyledonshave an indefinitenumberof floralparts in a low
spiral, while in the higher formsthe number is definiteand the
arrangementcyclic. The spur shoot of Pinus is deciduous in the
second to the twentiethyear (ELWES and HENRY 7, P. I002), and
its leaves fall with it, remainingpermanentlyattached. In the
othergenera it is the spur shoot whichis persistent,and the leaves
are cast either annually or in the second to the fifthyear. The
small and more or less definitenumberof persistentand whorled
needle leaves and the regularlydeciduous character of the spur
shoot are featureswhich render this structurein the pines very
unlike the ordinarybranch and also unlike the spurs of the other
forms,which differfroman ordinarybranch only in theirlimited
primary and secondary growth. The features that will receive
attention in this paper are such as indicate the branch character
of the spur in Pinus.
Number of needles
The number of leaves in the fasciclesof the pines, appearing
on firstsight constant,and being, as ENGLEMANN says (8), "the
most obvious distinctivecharacter,"has been extensivelymade use
of iji theirclassification. He states (8, p. i6i) that "the sections
of 2-leaved,3-leaved, and 5-leaved pines were the only ones known
to the older botanists"; to these were added two other sections
"by LINK (Linnaea, i84I), one with 2 or 3, the other with 3 or 4
leaves in a sheath." Subsequently formswith single leaves and
otherswith3-5 leaves in the fasciclehad to finda place. Numerous
exceptionsto this classificationhave also been recorded. KRON-
FELD (i6, p. 68) gives a summaryof the variationin many different
speciesobservedup to the date ofhis article. He cites,forexample,
the occurrencereportedby REICHARDT of 3, 4, and 5 needles in
P. silvestris,which is normallybifoliar,and also of 3, 4, and 6 in
364 BOTANICAL GAZETTE [MAY
Strobus. It is possible that they are P. excelsa,but, since some considerthe two
species as geographicalvarieties,the matteris unimportantfromthe presentstand-
point.
366 BOTANICAL GAZETTE [MAY
/
othertwo species,2but occurred
underthesame conditions.
/s 5 / ! In some youngplants of P.
ponderosa var. Jefreyi,I have
foundproliferationverycommon.
These plantswereabout 6 years
of age, and did not exhibitany
special vigor of growth. Text
fig. i shows the main axis and a
FIG. I PinUs ponderosa
var. Jef- branch,withsomeoftheneedles
freyi (one-halfnat. size): the main ofthespurfrom whichtheyarose
axis and a branch developedfrom attached to their basal regions.
spurshootsin a plantabout 6 years
old. Others of these needles have be-
comedetached. In theseplants
thereare normallyonly 3 needlesto a spur. In the axial stem-
producingspur shownabove,4 needlesare still attachedto the
base, and thebrokenstumpsof 2 otherscan be clearlyseen. The
branchspur has but 2 needlesintact,but the stumpsof 2 or 3
2Mr. C. H. MORSE, whileon a visitto the government nurseriesin Norfolk
County,was goodenoughto examineScotch,Banksian,and whitepine,and reports
is veryabundant,
that"proliferation especiallyin vigorous
plantsofP. silvestris
6 to
8 yearsofage,and fairlycommon
in P. BanksianaandP. Strobus ofaboutthesame
age.")
I9I4] THOMSON-SPUR SHOOT 379
has very long green bracts near its base and shows the gradation
between fascicled and primordialleaves, to which referencehas
alreadybeen made. I have also foundgreenbractson the branches
from spurs of P. excelsa, P. halepensis,P. Jefreyi,P. Laricio,
P. Pinaster, P. Pinea, P. Thunbergii,P. tuberculata. These are
especially large in the P. Thunbergii. I have also observed ordi-
nary proliferationdue to wounding in P. resinosa, P. silvestris,
and P. Strobus. In most cases these were young trees about io-
I5 feet high. In the case of P. Laricio, P. Laricio var. austriaca,
P. resinosa,P. silvestris,and P. Strobus,the trees were mature.
bifoliarspurs froman adult Scotch
In fig. I 2 two proliferating
pine are shown. It willbe noticedthat theyare the two uppermost
spurs (cf. DICKSON'S observationscited above), and that the injury
was done to the apex of the shoot of the year when the littlespurs
were just developing,not more than one-halfan inch in length
(see the one attached to the dead apex of the main axis). This is
a featurewhichis usual in otherspecies as well. When the injury
is done to the terminalpart of the young developingshoot, I have
always found a proliferationof the spurs below the injured part.
This never extends to the needles of a previous year, nor have I
observed that a spur shoot bud can ever be revived after it has
remaineddormantover the winter,unless some preliminarygrowth
took place the firstyear. This feature,however,is being investi-
gated by Mr. FRYER, who is carryingon a seriesof wound experi-
mentson the pines in order to determineat what season, on what
year's growth,etc., the best proliferationcan be obtainedby wound-
ing. He has not found that in the mature tree the spurs of P.
Strobus,when a year old, can be induced to develop, thoughthose
of the season's growthdo so very abundantly if wounded early.
His test cases were with twigs which had proliferatedlast year.
He tried to arouse the dormant buds of neighboringfasciclesby
cutting away all but the lowest and smallest of the proliferated
spurs. Only those that had grown the year before showed any
signsof doing so thisyear. It seems probable, then,in this species
that when the bud of the spur has lain dormant over the winter
it cannotbe revived. The leaves in thisformare shed in the second
year, and whetheror not those with more persistentspurs can be
I9I4] THOMSON-SPUR SHOOT 38i
LITERATURE CITED
i.BORTHWICK, A. W., On the developmentof quadrifoliarspurs in Pinus
Laricio Poir. Trans. and Proc. Bot. Soc. Edinburgh21:I50-I53. i899.
2. -, buds in pines. Ibid. I54-I58.
On interfoliar
3. BRITTON,N. L., NorthAmericanTrees. New York. i908.
4. COOLEY,GRACEE., Ecologicalnoteson the treesof theBotanical Gardens
at Naples. BOT. GAZ. 38:435-445. I904.
Morphologyof gymnosperms.Chicago.
5. COULTERand CHAMBERLAIN,
I9IO.
6. DICKSON,A., On the developmentof bifoliarspursinto ordinarybuds in
Pinus silvestris. Trans. and Proc. Bot. Soc. Edinburghi6:258-26i. i885.
7. ELWES,H. J., and HENRY, A., The treesof Great Britain and Ireland.
Edinburgh.i9io.
8. ENGELMANN, G., Revision of the genus Pinus. Trans. Acad. Sci. St.
Louis 4: i6i-i90. i88o.
W. M., The Potomac or youngerMesozoic flora. U.S. Geol.
9. FONTAINE,
Survey 15:I889.
IO. GOTHAN, W., Die fossilenHolzreste von Spitzbergen. Kungl. Svensk.
Vetensk.Akad. Handl. 45: no. 8. i9io.
i i. HEMPEL und WILHELM,Die Baume und Straucherdes Waldes. Wien.
I899.
12. HOCHSTETTER, W., Die sogenanntenRetinisporaArten der Garten.
Gartenfloravon E. REGEL. pp. 362-367. i88o.
13. of the Conif-
JEFFREY, E. C., The comparativeanatomyand phylogeny
erales. II. The Abietineae. Mem. Boston Soc. Nat. Hist. 6:I-37.
I905.
BOTANICAL GAZETTE, LVII PLATE XX
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THOMONoteSPRSHO
19I4] THOMSON-SPUR SHOOT 385
659. I904.
22. MEEHAN, T., Pinus edulis and P. monophylla.Bull. Torr. Bot. Club
ic2: 8i- 82. i885.
23. PHILLIPS, F. J., Two sprouting conifers of the Southwest. BOT. GAZ.
51:385-390. I9II.
24. SARGENT,C. S., The silva of NorthAmerica. Vol. XI. Boston. I897.
25. SHAw,G. R., Charactersof Pinus: the lateralcone. BOT. GAZ.43:205-
209. I907.
26. , The pines of Mexico. Publ. ArnoldArboretum,
no. i. Boston.
I909.
27. THOMSON, R. B., On the comparativeanatomyand affinities of the Arau-
carineae. Phil. Trans. Roy. Soc. London B 204:I-50. I9I3.
28. THOMSON,R. B., and ALLIN, A. E., Do the Abietineae extend to the Car-
boniferous? BOT. GAZ. 53:33I-338. I9I2.