Early  Journal  Content  on  JSTOR,  Free  to  Anyone  in  the  World  
This  article  is  one  of  nearly  500,000  scholarly  works  digitized  and  made  freely  available  to  everyone  in  
the  world  by  JSTOR.    

Known  as  the  Early  Journal  Content,  this  set  of  works  include  research  articles,  news,  letters,  and  other  
writings  published  in  more  than  200  of  the  oldest  leading  academic  journals.  The  works  date  from  the  
mid-­‐seventeenth  to  the  early  twentieth  centuries.    

 We  encourage  people  to  read  and  share  the  Early  Journal  Content  openly  and  to  tell  others  that  this  
resource  exists.    People  may  post  this  content  online  or  redistribute  in  any  way  for  non-­‐commercial  
purposes.  

Read  more  about  Early  Journal  Content  at  http://about.jstor.org/participate-­‐jstor/individuals/early-­‐

journal-­‐content.    

JSTOR  is  a  digital  library  of  academic  journals,  books,  and  primary  source  objects.  JSTOR  helps  people  
discover,  use,  and  build  upon  a  wide  range  of  content  through  a  powerful  research  and  teaching  
platform,  and  preserves  this  content  for  future  generations.  JSTOR  is  part  of  ITHAKA,  a  not-­‐for-­‐profit  
organization  that  also  includes  Ithaka  S+R  and  Portico.  For  more  information  about  JSTOR,  please  
contact  support@jstor.org.  
 ANATOMY OF LYCOPODIUM REFLEXUMI
J. BEN HILL

(WITH FIVE FIGURES)

In a formerpaper,2 giving in some detail the results of an

investigationof the development and specialization of the steles
of 6 species of Lycopodium,I reviewedthe literatureof the subject
of the anatomy of Lycopodium. The anatomical studies have
emphasized "types" of steles as characteristicof various species of
Lycopodium,among which the radial and parallel-banded arrange-
mentspredominate. In my formerpaper I suggestedthat it was
inadvisable to regardany stelar arrangementas characteristicof a
species of Lycopodium,since almost all "types" may be found
in a single species, and even in a single plant at differentlevels
in the stem.
The plants of L. reflexumused in this investigationwere col-
lected by Dr. C. R. BARNES and Dr. W. J. G. LAND in the vicinity
of Xalapa, Mexico, in i908. The habitat is described as moist
clay soil. The material was preservedin a formaldehyde-alcohol
solution and was given to me in this condition by Dr. LAND, to
whom I wish to express my thanks. The slides for the investi-
grationwere prepared fromparaffinserial sections cut transversely
10-I5 1Ain thickness and stainedin safranin-light greenand in
iron-alum haematoxylin-safranin,both combinations producing
excellent results; the formeris slightlybetter for differentiating
protoxylemin sections of young stems, and the latter better for
older tissues.
Investigation
In a study of the sections of the stem the most important
matter of interest is concerned with the so-called "types" of
stele to be foundin L. refexum; a secondarysignificantfeatureis
I Contributionsfrom the Department of Botany of the Pennsylvania State

College,no. I 7.
2 HILL, J. BEN, The anatomy of six epiphyticspecies of Lycopodium. BOT.

GAZ.58:6i-85. figs. 28. 1914.

BotanicalGazette,vol. 68] [2 26
19191 HILL-ANATOMY OF LYCOPODIUM 227

thepresenceofcorticalroots. Theseoriginatein a mannersimilar

to that describedby Miss STOKEY3for L. pithyoides,
and showa

S4~~~

4.--

4 ,

transversesectionof stem showingsmall stele and 3 cor-

FIG. xi.-L. reflex-umn:
tical roots; in the rootsphloem is surroundedby a crescent-shapedmass of xylem;
root stele is surroundedby very thin-walledcells which generallyare crushed and
brokenin sectioning;outercortexof rootcomposedof thick-walledcells; X 120.

development in thematterofdifferentiation
ofthesteleparalleling
that describedforthe stemof Lycopodium, in whichthe xylem
3 STOKEY, ALMA G., The rootsof Lycopodiumpitlzyoides.BOT. GAZ. 44:57-63.
pis. 5, 6. 1907.
4 HILL, J.BEN, 10c.cit.
228 BOTANICAL GAZETTE [SEPTEMBER

regionsare recognizablelongbeforesignification occurs. In young

sectionsof therootsthemetaxylem cellsare recognizableearlyby
theirincreasedsize and lack ofprotoplasmic content. The xylem
is crescent-shaped, the protoxylemoccurringat the ends and
bordering the largerarc of the crescent. The matureroots,2 or

FIG. 2.-L. reflexum: sectionthrough

transverse youngstemtipshowing several
protoxylem pointsalternating withphloemin stele; phloemextendstowardcenterof
cylinder, alternating withunlignifiedmetaxylem cells; thereis a phloemisland
entirelysurrounded by undignifiedmetaxylem cells; thisarrangementwhenmature
is represented in fig.5; X 200.

3 in a stem,are typicalcorticalroots(fig.i). The stemof Lyco-

podiumreflexum is small,about i mm.or less in diameter,witha
verysmallstele,0. 2 mm. in diameter.
In statingthe resultsof the investigation of the stelesof the
stemofL. reflexum I shallincludesimplya briefdescription ofthe
stelesfoundin the stemsof variousages, omittingthe detailsof
the developmentof the types,whichwere given in the former
articleand do notseemto varymuchin different species.
I9I9] IHILL-ANA TOMY OF LYCOPODIUM 229

radialarrangement
FIG. 3.-L. refiexurn:steleshowing of xylemwithphloem
located betweenprotoxylempointsand extendingtowardcenterof cylinder; X 200.

steleshowingparallel-banded
FIG. 4.-L. reflexum: arrangementof xylemwith
pbloem occurringin bands alternatingwith xylem and extendingacross cylinder;
X200.
230 BOTANICAL GAZETTE [SEPTEMBER

The numberof protoxylempoints in the stem stele ranges

from4 to ii, withabout 7 as themostfrequent number. Sections
of theyoungstemtip showtheconditioncharacteristic ofLycopo-
dium,in whichat about the timeof differentiation of the proto-
xylempoints the metaxylemcells are distinguishable by their
content
largesize and lack of protoplasmic (fig.2). There are 3
so-calledtypes of stele to be foundin L. reftexum: the radial
arrangement(fig. 3), the parallel-bandedarrangement(fig. 4),

innercylinder
steleshowing
FIG. 5.-L. reflexum: strand
of xylemsurrounding
ofphloem,mostfrequentarrangement X 200.
foundin steleofL. reflexum;

and an arrangement of an innercylinderof xylemsur-

consisting
roundinga strandof phloem(fig.5). These 3 arrangements of
the xylemmaybe foundin thesame stemat different levels,and
are all modificationsof the radial arrangement.The parallel-
banded arrangement, in whichalternatingstrandsof xylemand
phloemoccurin parallelbands acrossthe cylinder,seemsto be
correlatedto some extentwith the growthof roots,since this
arrangement is to be foundmostfrequently in the regionwhere
the rootsarise. The arrangement mostfrequently foundis that
consistingof an innercylinderof xylemenclosinga strandof
phloem(fig.5). Fromthis cylinderstrandsof xylemradiateto
1919] HIILL- NA TOMW
Y OF L X COPODIJUM.l 23 1

the protoxylem points. The condition is very similar to the

characteristicstele describedforL. Billardieri. The innercylinder
does not remain intact throughany great length of stem. but is
frequentlybroken up and gives rise to a parallel-banded arrange-
ment or revertsto the radial arrangement.

Summary
I. The 2 points of interest in the study of the anatomy of

Lycopodiumtreflexur are the presence of typical cortical roots

and the various "types" of stele in the stem.
2. The development and differentiation of the tissues in the
steles of the cortical roots parallel those in the stele of the stem.
3. There are 3 arrangementsof the xylem: radial, parallel-
banded, and a radial arrangementso modifiedas to consist of an
inner cylinderof xylem enclosinga small strand of phloem. The
last is the most frequentlyfound.
4. The study confirmsmy formersuggestionsthat all arrange-
ments of xylem may occur in the same stem in species of Lycopo-
dium.
THE PENNSYLVANIA STATE COLLEGE
STATE COLLEGE, PA.