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Research Article Morphometric and Statistical Analysis of the Palmaris Longus Muscle in Human and Non-Human Primates Roqueline A. G. M. F. Aversi-Ferreira, 1,2,3,4 Rafael Vieira Bretas, 5 Rafael Souto Maior, 3,4 Munkhzul Davaasuren, 5 Carlos Alberto Paraguassú-Chaves, 6 Hisao Nishijo, 5 and Tales Alexandre Aversi-Ferreira 1,2 1 Department of Anatomy, Howard University College of Medicine, 520 W Street NW, Numa Adams Building, Washington, DC 20059, USA 2 Laboratory of Primate Anthropology, Biochemistry, Neurosciences and Behavior, Federal University of Tocantins, NS 15 Avenue, Block 109 Norte, Plano Diretor Norte, 77001-090 Palmas, TO, Brazil 3 Graduate School of Animal Biology, Institute of Biology, University of Brasilia, Darcy Ribeiro Campus, 70910-900 Bras´ ılia, DF, Brazil 4 Department of Physiology, Laboratory of Neuroscience and Behavior, University of Brasilia, Darcy Ribeiro Campus, 70910-900 Bras´ ılia, DF, Brazil 5 System Emotional Science, Graduate School of Medicine and Pharmaceutical Sciences, University of Toyama, Toyama 930-1494, Japan 6 Federal University of Rondonia, RO, Brazil Correspondence should be addressed to Tales Alexandre Aversi-Ferreira; [email protected] Received 7 February 2014; Accepted 27 March 2014; Published 13 April 2014 Academic Editor: Carlos Tomaz Copyright © 2014 Roqueline A. G. M. F. Aversi-Ferreira et al. is is an open access article distributed under the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. e palmaris longus is considered a phylogenetic degenerate metacarpophalangeal joint flexor muscle in humans, a small vestigial forearm muscle; it is the most variable muscle in humans, showing variation in position, duplication, slips and could be reverted. It is frequently studied in papers about human anatomical variations in cadavers and in vivo, its variation has importance in medical clinic, surgery, radiological analysis, in studies about high-performance athletes, in genetics and anthropologic studies. Most studies about palmaris longus in humans are associated to frequency or case studies, but comparative anatomy in primates and comparative morphometry were not found in scientific literature. Comparative anatomy associated to morphometry of palmaris longus could explain the degeneration observed in this muscle in two of three of the great apes. Hypothetically, the comparison of the relative length of tendons and belly could indicate the pathway of the degeneration of this muscle, that is, the degeneration could be associated to increased tendon length and decreased belly from more primitive primates to those most derivate, that is, great apes to modern humans. In conclusion, in primates, the tendon of the palmaris longus increase from Lemuriformes to modern humans, that is, from arboreal to terrestrial primates and the muscle became weaker and tending to be missing. 1. Introduction e palmaris longus (PL) is considered a phylogenetic degen- erate metacarpophalangeal joint flexor muscle in humans [1] and a small vestigial forearm muscle [2]; it is the most variable muscle in humans [24], showing variation in position, duplication, and slips [2] and could be reverted [5]. It is frequently studied in papers about human anatomical variations in cadavers and in vivo [6]; its variation has importance in medical clinic, for example, vascular-neural compression [7], in surgery by using its tendon for graſting or other reconstructions [2, 4, 7], in radiological analysis, in study about high-performance athletes [8], and in genetics and anthropologic studies [2]. is muscle presents significant divergences as to its frequency in different humans groups [2, 7, 912]; it can be absent in some individuals of the genera Pan and Gorilla [6], Hindawi Publishing Corporation BioMed Research International Volume 2014, Article ID 178906, 6 pages http://dx.doi.org/10.1155/2014/178906

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Page 1: Research Article Morphometric and Statistical Analysis of ...downloads.hindawi.com/journals/bmri/2014/178906.pdf · frequency or case studies, but comparative anatomy in primates

Research ArticleMorphometric and Statistical Analysis of the Palmaris LongusMuscle in Human and Non-Human Primates

Roqueline A G M F Aversi-Ferreira1234 Rafael Vieira Bretas5

Rafael Souto Maior34 Munkhzul Davaasuren5 Carlos Alberto Paraguassuacute-Chaves6

Hisao Nishijo5 and Tales Alexandre Aversi-Ferreira12

1 Department of Anatomy Howard University College of Medicine 520 W Street NW Numa Adams BuildingWashington DC 20059 USA

2 Laboratory of Primate Anthropology Biochemistry Neurosciences and Behavior Federal University of TocantinsNS 15 Avenue Block 109 Norte Plano Diretor Norte 77001-090 Palmas TO Brazil

3 Graduate School of Animal Biology Institute of Biology University of Brasilia Darcy Ribeiro Campus 70910-900 Brasılia DF Brazil4Department of Physiology Laboratory of Neuroscience and Behavior University of Brasilia Darcy Ribeiro Campus70910-900 Brasılia DF Brazil

5 System Emotional Science Graduate School of Medicine and Pharmaceutical Sciences University of ToyamaToyama 930-1494 Japan

6 Federal University of Rondonia RO Brazil

Correspondence should be addressed to Tales Alexandre Aversi-Ferreira aversiferreiragmailcom

Received 7 February 2014 Accepted 27 March 2014 Published 13 April 2014

Academic Editor Carlos Tomaz

Copyright copy 2014 Roqueline A G M F Aversi-Ferreira et al This is an open access article distributed under the CreativeCommons Attribution License which permits unrestricted use distribution and reproduction in any medium provided theoriginal work is properly cited

The palmaris longus is considered a phylogenetic degenerate metacarpophalangeal joint flexor muscle in humans a small vestigialforearmmuscle it is the most variable muscle in humans showing variation in position duplication slips and could be reverted Itis frequently studied in papers about human anatomical variations in cadavers and in vivo its variation has importance in medicalclinic surgery radiological analysis in studies about high-performance athletes in genetics and anthropologic studiesMost studiesabout palmaris longus in humans are associated to frequency or case studies but comparative anatomy in primates and comparativemorphometry were not found in scientific literature Comparative anatomy associated to morphometry of palmaris longus couldexplain the degeneration observed in this muscle in two of three of the great apes Hypothetically the comparison of the relativelength of tendons and belly could indicate the pathway of the degeneration of this muscle that is the degeneration could beassociated to increased tendon length and decreased belly from more primitive primates to those most derivate that is great apesto modern humans In conclusion in primates the tendon of the palmaris longus increase from Lemuriformes to modern humansthat is from arboreal to terrestrial primates and the muscle became weaker and tending to be missing

1 Introduction

Thepalmaris longus (PL) is considered a phylogenetic degen-erate metacarpophalangeal joint flexor muscle in humans[1] and a small vestigial forearm muscle [2] it is the mostvariable muscle in humans [2ndash4] showing variation inposition duplication and slips [2] and could be reverted [5]It is frequently studied in papers about human anatomicalvariations in cadavers and in vivo [6] its variation has

importance in medical clinic for example vascular-neuralcompression [7] in surgery by using its tendon for graftingor other reconstructions [2 4 7] in radiological analysis instudy about high-performance athletes [8] and in geneticsand anthropologic studies [2]

This muscle presents significant divergences as to itsfrequency in different humans groups [2 7 9ndash12] it can beabsent in some individuals of the genera Pan and Gorilla [6]

Hindawi Publishing CorporationBioMed Research InternationalVolume 2014 Article ID 178906 6 pageshttpdxdoiorg1011552014178906

2 BioMed Research International

but it is always present inHylobatesPongo [9 10] and Sapajus[13] and its absence has not been reported in other primates

Most studies about PL in humans are associated withfrequency or case studies but comparative anatomy inprimates and comparative morphometry were not foundin scientific literature The comparative anatomy associatedto the morphometry of palmaris longus could explain thedegeneration observed in this muscle in two of three of thegreat apes (ie genera Pan andGorilla) andmodern humans

Hypothetically the comparison of the relative lengthof tendons and belly could indicate the pathway of thedegeneration of thismuscle that is the degeneration could beassociated with increased tendon length and decreased bellyfrom more primitive primates to those most derivate that isgreat apes to modern humans

Therefore the aim this work was to compare the anatomyand to verify the relative tendonbelly length of the palmarislongus in some primates from the NewWorld andOldWorldand modern humans associating data observed with thosefrom the literature

2 Material and Methods

Thiswork was previously approved by the Institutional EthicsCommittee of the Federal University of Goias state of GoiasBrazil (CoEP-UFG 812008) for human and nonhumanprimates studied in Brazil For other primates the rules ofanimal care in the USA and Japan were followed

21 Samples This study evaluated 14 adult human cadavers(males) allocated at the Laboratory of Human AnatomyFederal University of Goias Brazil six exemplars of adultSapajus libidinosus (5 males and 1 female) and one exemplarof adult Callithrix sp (male) allocated at the Laboratoryof Anthropology Biochemistry Neuroscience and PrimatesrsquoBehavior (LABINECOP) Federal University of TocantinsBrazil three exemplars of adult Macaca fuscata (males)allocated at the Laboratory of System Emotional ScienceToyama University Japan and one exemplar of adult Atelessp (male) one neonate Pongo sp (male) one neonate andone child Pan sp (males) one adult Callithrix sp (male)two Aotus sp two Lemur catta (one male one female) andone Propithecus sp (male) allocated at the Laboratory ofEvolutionary Biology Howard University USA

22 Dissection Documentation and Measures Humancadavers Sapajus sp and one forearm of child Pan sp hadbeen dissected for other purposes but other specimenswere dissected for purposes of this work The muscleswere photographed by digital camera and their lengthwas measured by metallic measure tape and digital politer(Niigara Seiki model DN 150) The values of measures werestandardized to specific measure of the digital politer Threemeasures of each structure were made and the average wasused for purpose of statistical analysis The tendons weremeasured from the point no part of the belly associatedto the tendon could be seen by naked eye to the palmaraponeurosis at the level of the radiorsquos head Some comparison

data from other primates were not observed in this workand were obtained from the literature for example Gorillasp and Hylobates sp The frequency of muscles wheneverpossible was based on scientific literature

23 Statistical Analyses Statistical analyses were performedusing the StatPlusmac AnalystSoft Inc2009 software tocalculate the average standard deviation and compare themean length values To effect comparative anatomy statisticswas performed based on Aversi-Ferreira [14] and Aversi-Ferreira et al [15] and data on muscle frequency origin andinsertion from literature [10 13] was used and comparedagainst data of this work To perform the statistics simplecomparative nonparametric method was used to comparetwo different species associated with anatomical conceptsof normality and variation as nominal variables Relativefrequency (RF) was defined as

RF = 119873 minus 119899V119873

(1)

where 119873 is the total number of specimens and 119899V is thenumber of individuals presenting variation of the normalpattern therefore RF means the structure normality in apopulation sample

When more than one parameter was used they wererelated to specific weighted values with respect to their degreeof relevance in comparative analysis Parameters with lessvariation in phylogenetic terms were assigned a higher valueTherefore origin insertion innervation and presence ofmuscle in terms of frequency and type of fiber arrangementwere assigned weights 4 4 3 2 and 1 respectively

The weighted average of frequencies (WAF) was calcu-lated using the RF values

PAF

=

RF1times 1198751+ RF2times 1198752+ RF3times 1198753+ RF4times 1198754+ RF5times 1198755

1198751+ 1198752+ 1198753+ 1198754+ 1198755

(2)

where RF1is the frequency of musclersquos origin and 119875

1is 4 RF

2

is the frequency of musclersquos insertion and 1198752is 4 RF

3is the

frequency of innervation and 1198753is 3 RF

4is the frequency of

the presence of muscle and 1198754is 2 and RF

5is the type of

belly arrangement and the 1198755is 1 The frequencies of RF to

humans Pan sp and Gorilla sp were obtained from Gibbs[10] who cited the absence of palmaris longus from 39 to204 therefore 119899V was calculated based on the average ofthese numbers that is 1215 and 119873 was considered to be100 therefore RF to human was 0878 The 119899V value forchimpanzee was 19 and the119873 value was 28 and RFwas 0678for gorilla the 119899V value was 6 and the119873 value was 19 and RFwas 0316 The type of fiber arrangement was considered tobe 1 to Lemuriformes and NewWorld primates and 05 to theothers because this was considered here as an intermediatecharacter among these primates

To consider the phylogenetic proximity between thestructures studied the difference in the relative frequencywas

BioMed Research International 3

(a) (b) (c) (d) (e) (f)

(g) (h) (i)

Figure 1 Photos of the forearms of (a) Propithecus sp (045x left) (b) Lemur catta (052x left) (c) Sapajus libidinosus (034x right) (d)Atelessp (01x right) (e) Callithrix sp (03x right) (f) Aotus sp (08x right) (g) Macaca fuscata (034x right) (h) Pongo sp (079x left) (i) Pansp (023x left) From (a) to (f) muscles are pennate and from (g) to (i) are fusiform lowast indicates the palmaris longus

calculated or Comparative Anatomy Index (CAI) betweensamples from different species

CAI = 1003816100381610038161003816

PAF119894minus PAF

119894119894

1003816100381610038161003816

(3)

where indices 119894 and 119894119894 represent samples 1 and 2From the previous equations it follows that CAI value

close to zero represents greater similarity between sam-ples whereas CAI closer to 10 implies higher divergencebetween samples Therefore the greater the numerical dis-tance between values the greater the divergence in relationto the palmaris longus muscle between species

3 Results

In all primates studied here the PL tendon originated frommedial humeral epicondyle and inserted into the palmar

fascia was innervated by the median nerve A close relation-ship between the palmaris longus tendon and the fascia ofthe forearm was observed which is similar to other flexormuscles of the forearm The belly of palmaris longus waseasily distinguished from the other muscles of the forearmin all studied species except for the only exemplar of Atelessp in which the flexor muscles formed one group of belliesfrom the elbowwith a separation of tendons close to the wrist(Figure 1)

Regarding the type of fiber arrangement in the belly thepalmaris longus tendon presented an aspect similar to pen-nate in all Lemuriformes and NewWorld primates (Figure 1)but fusiform to Macaca fuscata (unique exemplar of OldWorld primates) and to apes (Pongo sp and Pan sp) andmodern humansThe average palmaris longustendon lengthshowed significant differences between species studied in thiswork (Table 1 Figure 2) Shorter tendons were observed in

4 BioMed Research International

Table 1 Measures of the palmaris longus palmaris longus tendon and palmaris longustendon relationship for species of primates andprimate groups

Specimens (119899) Total Tendon Totaltendon Primate groupsModern humans (28) 2551 (plusmn169) 1506 (plusmn175) 171 (plusmn013) Modern humansdaggerΔlowast+

Pan sp (3) 883 (plusmn162) 495 (plusmn091) 178 (plusmn004) ApesdaggerΔlowast+Pongo sp (2) 830 (plusmn010) 440 (plusmn030) 189 (plusmn015)Macaca fuscata (6) 1730 (plusmn043) 730 (plusmn05) 237 (plusmn012) Old World primatesdaggerΔlowast

Aotus sp (4) 68 (plusmn047) 32 (plusmn050) 209 (plusmn023)

NewWorld primatesdaggerΔCallithrix sp (3) 435 (plusmn019) 172 (plusmn023) 253 (plusmn008)Ateles sp (1) 253 100 253Sapajus libidinosus (11) 1071 (plusmn150) 299 (plusmn051) 381 (plusmn107)Lemur catta (2) 86 (plusmn050) 19 (plusmn00) 453 (plusmn027) LemuriformesdaggerPropithecus sp (2) 1125 (plusmn005) 22 (plusmn020) 516 (plusmn049)daggerSignificant difference among Lemuriformes and other groupsΔSignificant difference among NewWorld primates and other groupslowast Significant difference among Old World primates and other groups+Significant difference between apes and modern humans

Table 2 Statistics of the comparative anatomy of palmaris longus

Taxon PAF PAF =RF1 times 1198751 + RF2 times 1198752 + RF3 times 1198753 + RF4 times 1198754 + RF5 times 1198755

1198751+ 1198752+ 1198753+ 1198754+ 1198755

CAI = |PAF119894minus PAF

119894119894|

Propithecus sp 1 PAF = 4 times 1 + 4 times 1 + 3 times 1 + 2 times 1 + 1 times 114

Reference (most primitive characters)

Lemur catta 1 PAF = 4 times 1 + 4 times 1 + 3 times 1 + 2 times 1 + 1 times 114

0

Callithrix sp 1 PAF = 4 times 1 + 4 times 1 + 3 times 1 + 2 times 1 + 1 times 114

0

Sapajus sp 1 PAF = 4 times 1 + 4 times 1 + 3 times 1 + 2 times 1 + 1 times 114

0

Ateles sp 1 PAF = 4 times 1 + 4 times 1 + 3 times 1 + 2 times 1 + 1 times 114

0

Macaca fuscata 0964 PAF = 4 times 1 + 4 times 1 + 3 times 1 + 2 times 1 + 1 times 0514

0036

Pongo sp 0964 PAF = 4 times 1 + 4 times 1 + 3 times 1 + 2 times 1 + 1 times 0514

0036

Modern humans 0947 PAF = 4 times 1 + 4 times 1 + 3 times 1 + 2 times 0878 + 1 times 0514

0053

Pan sp 0918 PAF = 4 times 1 + 4 times 1 + 3 times 1 + 2 times 0678 + 1 times 0514

0082

Gorilla sp 0867 PAF = 4 times 1 + 4 times 1 + 3 times 1 + 2 times 0316 + 1 times 0514

0133

PAF is the weighted average of frequencies CAI is Comparative Anatomy Index

Lemuriformes and longer in modern humans (119875 lt 005Figure 2)

Significant differences were observed between averages(119875 lt 005) calculated for all groups that is Lemuriformes andother groups NewWorld primates and other groups andOldWorld primates (Macaca fuscata) and other groups and forapes andmodern humans (Table 1 Figure 2) Nevertheless toAotus sp specifically in comparison to Macaca fuscata andapes differences were not observed

The CAI indicates that Lemuriformes and New Worldprimates share similar characters to PL (CAI = 0) while

these features are more derived in Gorilla sp (CAI = 0133)(Table 2)

4 Discussion

The large variations in the prevalence of the PL amongmodern humans may be indicative that this muscle is degen-erating [1] and its small belly may suggest it is a vestigialmuscle [2]

Although there are several studies investigating the fre-quency of PL in modern humans only a few recent studies

BioMed Research International 5

Lemuriformes New Worldprimates primates

Old World Apes Modernhumans

BoxMean line

0

1

2

3

4

5

6

Palm

aris

long

uste

ndon

(cm

)

Figure 2 Graph showing the mean line relative to the palmarislongustendon length of primate groups

have compared PL frequency in nonhuman primates [9 1016ndash18] From such studies it was found for example that thefrequency of PL in Gorilla sp is much lower than in humansand chimpanzees even though humans and chimpanzees arethe only species in which absence of PL has been reportedThe PL is found in Gorilla sp from 15 to 63 accordingto some authors [10 16 17 19 20] and could be consideredabsent in gorillas according to [18]

In this sense considering only the frequency of thismuscle in primates PLwould bemore degenerative inGorillasp Also if degeneration is considered a derived characteristicof PL it is more derived in Gorilla sp than in other primatesand modern humans Indeed it is reasonable to suggest thatPL is a degenerative muscle because according to Meglioli[21] its agenesis is associated with a recessive gene

Notwithstanding when parameters such as origin inser-tion innervation frequency and type of belly fibers arecompared together and using specific weight (ie whenCAI is applied for review see Aversi-Ferreira [14] andAversi-Ferreira et al [15]) Gorilla sp shows the greatestnumerical distance from the primitive character consideredin Propithecus sp followed by Pan sp modern humansPongo sp and Macaca fuscata in decreasing order

In fact the different variation factor observed in the CAIcalculation can be explained by the low frequency in thenumber of specimens and species and by the different types ofmuscle fibers of the belly muscle obtained from the literatureand from the present study To Lemuriformes andNewWorldprimates CAI indicates no quantitative difference that is thecalculated value is zero but toMacaca fuscata and Pongo spthe only difference is type of fiber arrangement

In order to obtain more objective parameters the relativelength of the tendon was studied that is the length of thePL muscle divided by the length of the palmaris longus

tendon Its measure indicates indirectly the muscle relativestrength because smaller belly indicates less sarcomeresacting together to generate contraction [22] Therefore alonger tendon indicates less muscular force According tothe data obtained here this relation (PL muscle divided bythe length of the palmaris longus tendon) decreases fromLemuriformes to modern humans

Interestingly the comparison between means of therelative measures of PL among these groups of primates(119875 lt 005) showed significant differences between one groupand all other groups One discrepancy was observed in themeasures obtained from Aotus sp a New World primate inwhich the average measures were similar to Macaca fuscataand apes but it was not possible to associate it with any aspectconsidered in this work Putatively a weak PL is observedin modern humans and apes and strong in LemuriformesTherefore it is possible to hypothesize that the tendon lengthdecreases from arboreal to terrestrial primates In line withthis Ankel-simons [23] reports that great apes move in aquadrupedal Knuckle-walking manner the Macaca genusis terrestrial quadruped all New World primates are highlyarboreal and all Lemuriformes except for Lemur catta spendaround 13 of the time on the ground

Specifically to modern humans the average measure ofthe PL tendon obtained here that is 1506 (plusmn175) wasdifferent from those reported for other Brazilians cadavers[12] (1199 plusmn 152) Apparently these differences are due tocriteria for length measurements of the palmaris longustendon in both studies

On the other hand different fiber arrangement of thebelly of PL in Lemuriformes and New World primatescompared to OldWorld primates apes and modern humanswas observed The tendon begins in the belly which involvesthe tendon laterally up to approximately the wrist in Lemu-riformes and New World primates characterizing a pennatemuscle whereas fromMacaca fuscata tomodern humans thetendon begins after the end of the belly observed via nakedeye characterizing a fusiform muscle

This characteristic affects the muscular strength thefusiform muscle generates a more direct contraction butto modern humans apes and Macaca fuscata the belliesare shorter in a pennate muscle however the physiologicalcross-sectional area is considerably larger than its anatomicalcross-sectional area Therefore the pennate muscle ceterisparibus generates more force [24] In addition they arelonger in Lemuriformes and New World primate than inother primates

The number of animals per species and the number ofspecies used here allowed inferring the following conclusionsFirst the more derived characteristics of PL are found inapes especially in Gorilla sp relative to evolutionary aspectsmeasured by CAI but a shorter tendon is verified in modernhumans Second the path to the degeneration of the PL seemsto follow a decreased tendon associated with modification inthe type of belly fibers from pennate to fusiform therefore inprimates with more derived thoracic limbs the weaker PL

To our knowledge this is the first study to present anevolutionary anatomical comparison of the PL Further stud-ies could indicate more accurately the evolutionary pathway

6 BioMed Research International

followed by PL from primitive to derived primates associatedwith aspects such as the muscular force difference betweenprimates and verification of anatomical aspects in morespecies and consider intra- and interspecific variations

5 Conclusions

Apparently in primates the PL tendon relative size seems toincrease from ancestors genera to more derived ones Thissuggests that this muscle is weaker in terrestrial primateswhen compared to arboreal primates The PL tendon seemsto be more derived in Gorilla sp considering the factorsanalyzed here but the tendon is longer in modern humans

Conflict of Interests

The authors declare that there is no conflict of interestsregarding the publication of this paper

References

[1] S Standring ldquoPelvis girdle and lower limbrdquo in Grayrsquos Anatomythe Anatomical Basis of Clinical Practice Churchill LivingstoneLondon UK 2008

[2] J W M Kigera and S Mukwaya ldquoFrequency of agenesisPalmaris longus through clinical examinationmdashan East Africanstudyrdquo PLoS ONE vol 6 no 12 Article ID e28997 2011

[3] H Gray ldquoMembro superiorrdquo in Gray Anatomia GuanabaraKoogan Rio de Janeiro Brazil 1979

[4] V S Telles and B A L Ernesto ldquoConsideraciones anatomicasde los musculos inconstantesrdquo MedUnab vol 1 no 3 pp 165ndash170 1998

[5] J M Cope E M Looney C A Craig R Gawron R Lamprosand R Mahoney ldquoMedian nerve compression and the reversedpalmaris longusrdquo International Journal od Anatomical Varia-tion vol 2 pp 102ndash104 2009

[6] A B M Machado and L J A Di Dio ldquoFrequency of themusculus palmaris longus studied in vivo in some Amazonindiansrdquo American Journal of Physical Anthropology vol 27 no1 pp 11ndash20 1967

[7] M A Babinski K M A Valente C M S Savedra C H FBurity and A M S Sabra ldquoVariacao anatomica do musculopalmar longo associado a ausencia do arco palmar superficiale compressao do canal de GuyonrdquoActa Scienteae Medica vol 2no 2 pp 54ndash58 2008

[8] C Fowlie C Fuller and M K Pratten ldquoAssessment of thepresenceabsence of the palmaris longus muscle in differentsports and elite andnon-elite sport populationsrdquoPhysiotherapyvol 98 no 2 pp 138ndash142 2012

[9] L Testut and A Latarjet Tratado de Anatomia Humana SalvatEditores Barcelona Espana 1959

[10] S Gibbs Comparative soft tissue morphology of the extantHominoidea including man [PhD thesis] The University ofLiverpool Inglaterra 1999

[11] A O Kayode A A Olamide I O Blessing and O U VictorldquoIncidence of palmaris longus muscle absence in Nigerianpopulationrdquo International Journal of Morphology vol 26 no 2pp 305ndash308 2008

[12] L C Angelini Junior F B Angelini B C Oliveira S ASoares L C Angelini and R H Cabral ldquoUse of the tendon

of the palmaris longus muscle in surgical proedures study oncadaversrdquoActaOrtopedica Brasileira vol 20 no 4 pp 226ndash2292012

[13] T A Aversi-Ferreira L G Vieira R M Pires Z Silvaand N Penha-Silva ldquoEstudo anatomico dos musculos flexoressuperficiais do antebraco no macaco Cebus apellardquo BioscienceJournal vol 22 no 1 pp 139ndash144 2006

[14] T A Aversi-Ferreira ldquoA new statistical method for comparativeanatomyrdquo International Journal of Morphology vol 27 no 4 pp1051ndash1058 2009

[15] T A Aversi-Ferreira R S Maior F O Carneiro-e-Silva et alldquoComparative anatomical analyses of the forearm muscles ofCebus libidinosus (Rylands et al 2000) manipulatory behaviorand tool userdquo PLoS ONE vol 6 no 7 Article ID e22165 8 pages2011

[16] H Preuschoft ldquoMuscles and joints of the anterior extremity ofa gorilla (Gorilla gorilla Savage et Wyman 1847)rdquo Gegenbaursmorphologisches Jahrbuch vol 107 no 2 pp 99ndash183 1965

[17] E E Sarmiento ldquoTerrestrial traits in the hands and feet ofgorillasrdquo American Museum Novitates vol 3091 pp 1ndash56

[18] R Diogo J F Pastor E M Ferrero et al Photographic andDescriptive Musculoskeletal Atlas of Gorilla With Notes on theAttachments Veriations Innervation Synonymy and Weight ofthe Muscles Science Publisher Enfield USA 2010

[19] A Keith ldquoOn the chimpanzees and their relationship to thegorillardquo Proceedings of the Zoological Society of London vol 67no 2 pp 296ndash312 1899

[20] E Loth Anthropologie Des Parties Molles (Muscles IntestinsVaisseaux Nerfs Peripheriques) Mianowski-Masson et CieParis France 1931

[21] G T Meglioli ldquoEtude de Irsquoheredite du muscle petit palmaire(Musculus palmaris longus) dans la population belgerdquo BullEtinde La Societe Royale Belge DrsquoAnthropologie et de Prehistoire vol75 pp 67ndash86 1965

[22] PW Brand R B Beach and D EThompson ldquoRelative tensionand potential excursion of muscles in the forearm and handrdquoJournal of Hand Surgery vol 6 no 3 pp 210ndash219 1981

[23] F Ankel-Simons Primate Anatomy An Introduction AcademicPress Orlando Fla USA 2000

[24] Y Ichinose H KanehisaM Ito Y Kawakami and T FukunagaldquoRelationship between muscle fiber pennation and force gen-eration capability in Olympic athletesrdquo International Journal ofSports Medicine vol 19 no 8 pp 541ndash546 1998

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Microbiology

Page 2: Research Article Morphometric and Statistical Analysis of ...downloads.hindawi.com/journals/bmri/2014/178906.pdf · frequency or case studies, but comparative anatomy in primates

2 BioMed Research International

but it is always present inHylobatesPongo [9 10] and Sapajus[13] and its absence has not been reported in other primates

Most studies about PL in humans are associated withfrequency or case studies but comparative anatomy inprimates and comparative morphometry were not foundin scientific literature The comparative anatomy associatedto the morphometry of palmaris longus could explain thedegeneration observed in this muscle in two of three of thegreat apes (ie genera Pan andGorilla) andmodern humans

Hypothetically the comparison of the relative lengthof tendons and belly could indicate the pathway of thedegeneration of thismuscle that is the degeneration could beassociated with increased tendon length and decreased bellyfrom more primitive primates to those most derivate that isgreat apes to modern humans

Therefore the aim this work was to compare the anatomyand to verify the relative tendonbelly length of the palmarislongus in some primates from the NewWorld andOldWorldand modern humans associating data observed with thosefrom the literature

2 Material and Methods

Thiswork was previously approved by the Institutional EthicsCommittee of the Federal University of Goias state of GoiasBrazil (CoEP-UFG 812008) for human and nonhumanprimates studied in Brazil For other primates the rules ofanimal care in the USA and Japan were followed

21 Samples This study evaluated 14 adult human cadavers(males) allocated at the Laboratory of Human AnatomyFederal University of Goias Brazil six exemplars of adultSapajus libidinosus (5 males and 1 female) and one exemplarof adult Callithrix sp (male) allocated at the Laboratoryof Anthropology Biochemistry Neuroscience and PrimatesrsquoBehavior (LABINECOP) Federal University of TocantinsBrazil three exemplars of adult Macaca fuscata (males)allocated at the Laboratory of System Emotional ScienceToyama University Japan and one exemplar of adult Atelessp (male) one neonate Pongo sp (male) one neonate andone child Pan sp (males) one adult Callithrix sp (male)two Aotus sp two Lemur catta (one male one female) andone Propithecus sp (male) allocated at the Laboratory ofEvolutionary Biology Howard University USA

22 Dissection Documentation and Measures Humancadavers Sapajus sp and one forearm of child Pan sp hadbeen dissected for other purposes but other specimenswere dissected for purposes of this work The muscleswere photographed by digital camera and their lengthwas measured by metallic measure tape and digital politer(Niigara Seiki model DN 150) The values of measures werestandardized to specific measure of the digital politer Threemeasures of each structure were made and the average wasused for purpose of statistical analysis The tendons weremeasured from the point no part of the belly associatedto the tendon could be seen by naked eye to the palmaraponeurosis at the level of the radiorsquos head Some comparison

data from other primates were not observed in this workand were obtained from the literature for example Gorillasp and Hylobates sp The frequency of muscles wheneverpossible was based on scientific literature

23 Statistical Analyses Statistical analyses were performedusing the StatPlusmac AnalystSoft Inc2009 software tocalculate the average standard deviation and compare themean length values To effect comparative anatomy statisticswas performed based on Aversi-Ferreira [14] and Aversi-Ferreira et al [15] and data on muscle frequency origin andinsertion from literature [10 13] was used and comparedagainst data of this work To perform the statistics simplecomparative nonparametric method was used to comparetwo different species associated with anatomical conceptsof normality and variation as nominal variables Relativefrequency (RF) was defined as

RF = 119873 minus 119899V119873

(1)

where 119873 is the total number of specimens and 119899V is thenumber of individuals presenting variation of the normalpattern therefore RF means the structure normality in apopulation sample

When more than one parameter was used they wererelated to specific weighted values with respect to their degreeof relevance in comparative analysis Parameters with lessvariation in phylogenetic terms were assigned a higher valueTherefore origin insertion innervation and presence ofmuscle in terms of frequency and type of fiber arrangementwere assigned weights 4 4 3 2 and 1 respectively

The weighted average of frequencies (WAF) was calcu-lated using the RF values

PAF

=

RF1times 1198751+ RF2times 1198752+ RF3times 1198753+ RF4times 1198754+ RF5times 1198755

1198751+ 1198752+ 1198753+ 1198754+ 1198755

(2)

where RF1is the frequency of musclersquos origin and 119875

1is 4 RF

2

is the frequency of musclersquos insertion and 1198752is 4 RF

3is the

frequency of innervation and 1198753is 3 RF

4is the frequency of

the presence of muscle and 1198754is 2 and RF

5is the type of

belly arrangement and the 1198755is 1 The frequencies of RF to

humans Pan sp and Gorilla sp were obtained from Gibbs[10] who cited the absence of palmaris longus from 39 to204 therefore 119899V was calculated based on the average ofthese numbers that is 1215 and 119873 was considered to be100 therefore RF to human was 0878 The 119899V value forchimpanzee was 19 and the119873 value was 28 and RFwas 0678for gorilla the 119899V value was 6 and the119873 value was 19 and RFwas 0316 The type of fiber arrangement was considered tobe 1 to Lemuriformes and NewWorld primates and 05 to theothers because this was considered here as an intermediatecharacter among these primates

To consider the phylogenetic proximity between thestructures studied the difference in the relative frequencywas

BioMed Research International 3

(a) (b) (c) (d) (e) (f)

(g) (h) (i)

Figure 1 Photos of the forearms of (a) Propithecus sp (045x left) (b) Lemur catta (052x left) (c) Sapajus libidinosus (034x right) (d)Atelessp (01x right) (e) Callithrix sp (03x right) (f) Aotus sp (08x right) (g) Macaca fuscata (034x right) (h) Pongo sp (079x left) (i) Pansp (023x left) From (a) to (f) muscles are pennate and from (g) to (i) are fusiform lowast indicates the palmaris longus

calculated or Comparative Anatomy Index (CAI) betweensamples from different species

CAI = 1003816100381610038161003816

PAF119894minus PAF

119894119894

1003816100381610038161003816

(3)

where indices 119894 and 119894119894 represent samples 1 and 2From the previous equations it follows that CAI value

close to zero represents greater similarity between sam-ples whereas CAI closer to 10 implies higher divergencebetween samples Therefore the greater the numerical dis-tance between values the greater the divergence in relationto the palmaris longus muscle between species

3 Results

In all primates studied here the PL tendon originated frommedial humeral epicondyle and inserted into the palmar

fascia was innervated by the median nerve A close relation-ship between the palmaris longus tendon and the fascia ofthe forearm was observed which is similar to other flexormuscles of the forearm The belly of palmaris longus waseasily distinguished from the other muscles of the forearmin all studied species except for the only exemplar of Atelessp in which the flexor muscles formed one group of belliesfrom the elbowwith a separation of tendons close to the wrist(Figure 1)

Regarding the type of fiber arrangement in the belly thepalmaris longus tendon presented an aspect similar to pen-nate in all Lemuriformes and NewWorld primates (Figure 1)but fusiform to Macaca fuscata (unique exemplar of OldWorld primates) and to apes (Pongo sp and Pan sp) andmodern humansThe average palmaris longustendon lengthshowed significant differences between species studied in thiswork (Table 1 Figure 2) Shorter tendons were observed in

4 BioMed Research International

Table 1 Measures of the palmaris longus palmaris longus tendon and palmaris longustendon relationship for species of primates andprimate groups

Specimens (119899) Total Tendon Totaltendon Primate groupsModern humans (28) 2551 (plusmn169) 1506 (plusmn175) 171 (plusmn013) Modern humansdaggerΔlowast+

Pan sp (3) 883 (plusmn162) 495 (plusmn091) 178 (plusmn004) ApesdaggerΔlowast+Pongo sp (2) 830 (plusmn010) 440 (plusmn030) 189 (plusmn015)Macaca fuscata (6) 1730 (plusmn043) 730 (plusmn05) 237 (plusmn012) Old World primatesdaggerΔlowast

Aotus sp (4) 68 (plusmn047) 32 (plusmn050) 209 (plusmn023)

NewWorld primatesdaggerΔCallithrix sp (3) 435 (plusmn019) 172 (plusmn023) 253 (plusmn008)Ateles sp (1) 253 100 253Sapajus libidinosus (11) 1071 (plusmn150) 299 (plusmn051) 381 (plusmn107)Lemur catta (2) 86 (plusmn050) 19 (plusmn00) 453 (plusmn027) LemuriformesdaggerPropithecus sp (2) 1125 (plusmn005) 22 (plusmn020) 516 (plusmn049)daggerSignificant difference among Lemuriformes and other groupsΔSignificant difference among NewWorld primates and other groupslowast Significant difference among Old World primates and other groups+Significant difference between apes and modern humans

Table 2 Statistics of the comparative anatomy of palmaris longus

Taxon PAF PAF =RF1 times 1198751 + RF2 times 1198752 + RF3 times 1198753 + RF4 times 1198754 + RF5 times 1198755

1198751+ 1198752+ 1198753+ 1198754+ 1198755

CAI = |PAF119894minus PAF

119894119894|

Propithecus sp 1 PAF = 4 times 1 + 4 times 1 + 3 times 1 + 2 times 1 + 1 times 114

Reference (most primitive characters)

Lemur catta 1 PAF = 4 times 1 + 4 times 1 + 3 times 1 + 2 times 1 + 1 times 114

0

Callithrix sp 1 PAF = 4 times 1 + 4 times 1 + 3 times 1 + 2 times 1 + 1 times 114

0

Sapajus sp 1 PAF = 4 times 1 + 4 times 1 + 3 times 1 + 2 times 1 + 1 times 114

0

Ateles sp 1 PAF = 4 times 1 + 4 times 1 + 3 times 1 + 2 times 1 + 1 times 114

0

Macaca fuscata 0964 PAF = 4 times 1 + 4 times 1 + 3 times 1 + 2 times 1 + 1 times 0514

0036

Pongo sp 0964 PAF = 4 times 1 + 4 times 1 + 3 times 1 + 2 times 1 + 1 times 0514

0036

Modern humans 0947 PAF = 4 times 1 + 4 times 1 + 3 times 1 + 2 times 0878 + 1 times 0514

0053

Pan sp 0918 PAF = 4 times 1 + 4 times 1 + 3 times 1 + 2 times 0678 + 1 times 0514

0082

Gorilla sp 0867 PAF = 4 times 1 + 4 times 1 + 3 times 1 + 2 times 0316 + 1 times 0514

0133

PAF is the weighted average of frequencies CAI is Comparative Anatomy Index

Lemuriformes and longer in modern humans (119875 lt 005Figure 2)

Significant differences were observed between averages(119875 lt 005) calculated for all groups that is Lemuriformes andother groups NewWorld primates and other groups andOldWorld primates (Macaca fuscata) and other groups and forapes andmodern humans (Table 1 Figure 2) Nevertheless toAotus sp specifically in comparison to Macaca fuscata andapes differences were not observed

The CAI indicates that Lemuriformes and New Worldprimates share similar characters to PL (CAI = 0) while

these features are more derived in Gorilla sp (CAI = 0133)(Table 2)

4 Discussion

The large variations in the prevalence of the PL amongmodern humans may be indicative that this muscle is degen-erating [1] and its small belly may suggest it is a vestigialmuscle [2]

Although there are several studies investigating the fre-quency of PL in modern humans only a few recent studies

BioMed Research International 5

Lemuriformes New Worldprimates primates

Old World Apes Modernhumans

BoxMean line

0

1

2

3

4

5

6

Palm

aris

long

uste

ndon

(cm

)

Figure 2 Graph showing the mean line relative to the palmarislongustendon length of primate groups

have compared PL frequency in nonhuman primates [9 1016ndash18] From such studies it was found for example that thefrequency of PL in Gorilla sp is much lower than in humansand chimpanzees even though humans and chimpanzees arethe only species in which absence of PL has been reportedThe PL is found in Gorilla sp from 15 to 63 accordingto some authors [10 16 17 19 20] and could be consideredabsent in gorillas according to [18]

In this sense considering only the frequency of thismuscle in primates PLwould bemore degenerative inGorillasp Also if degeneration is considered a derived characteristicof PL it is more derived in Gorilla sp than in other primatesand modern humans Indeed it is reasonable to suggest thatPL is a degenerative muscle because according to Meglioli[21] its agenesis is associated with a recessive gene

Notwithstanding when parameters such as origin inser-tion innervation frequency and type of belly fibers arecompared together and using specific weight (ie whenCAI is applied for review see Aversi-Ferreira [14] andAversi-Ferreira et al [15]) Gorilla sp shows the greatestnumerical distance from the primitive character consideredin Propithecus sp followed by Pan sp modern humansPongo sp and Macaca fuscata in decreasing order

In fact the different variation factor observed in the CAIcalculation can be explained by the low frequency in thenumber of specimens and species and by the different types ofmuscle fibers of the belly muscle obtained from the literatureand from the present study To Lemuriformes andNewWorldprimates CAI indicates no quantitative difference that is thecalculated value is zero but toMacaca fuscata and Pongo spthe only difference is type of fiber arrangement

In order to obtain more objective parameters the relativelength of the tendon was studied that is the length of thePL muscle divided by the length of the palmaris longus

tendon Its measure indicates indirectly the muscle relativestrength because smaller belly indicates less sarcomeresacting together to generate contraction [22] Therefore alonger tendon indicates less muscular force According tothe data obtained here this relation (PL muscle divided bythe length of the palmaris longus tendon) decreases fromLemuriformes to modern humans

Interestingly the comparison between means of therelative measures of PL among these groups of primates(119875 lt 005) showed significant differences between one groupand all other groups One discrepancy was observed in themeasures obtained from Aotus sp a New World primate inwhich the average measures were similar to Macaca fuscataand apes but it was not possible to associate it with any aspectconsidered in this work Putatively a weak PL is observedin modern humans and apes and strong in LemuriformesTherefore it is possible to hypothesize that the tendon lengthdecreases from arboreal to terrestrial primates In line withthis Ankel-simons [23] reports that great apes move in aquadrupedal Knuckle-walking manner the Macaca genusis terrestrial quadruped all New World primates are highlyarboreal and all Lemuriformes except for Lemur catta spendaround 13 of the time on the ground

Specifically to modern humans the average measure ofthe PL tendon obtained here that is 1506 (plusmn175) wasdifferent from those reported for other Brazilians cadavers[12] (1199 plusmn 152) Apparently these differences are due tocriteria for length measurements of the palmaris longustendon in both studies

On the other hand different fiber arrangement of thebelly of PL in Lemuriformes and New World primatescompared to OldWorld primates apes and modern humanswas observed The tendon begins in the belly which involvesthe tendon laterally up to approximately the wrist in Lemu-riformes and New World primates characterizing a pennatemuscle whereas fromMacaca fuscata tomodern humans thetendon begins after the end of the belly observed via nakedeye characterizing a fusiform muscle

This characteristic affects the muscular strength thefusiform muscle generates a more direct contraction butto modern humans apes and Macaca fuscata the belliesare shorter in a pennate muscle however the physiologicalcross-sectional area is considerably larger than its anatomicalcross-sectional area Therefore the pennate muscle ceterisparibus generates more force [24] In addition they arelonger in Lemuriformes and New World primate than inother primates

The number of animals per species and the number ofspecies used here allowed inferring the following conclusionsFirst the more derived characteristics of PL are found inapes especially in Gorilla sp relative to evolutionary aspectsmeasured by CAI but a shorter tendon is verified in modernhumans Second the path to the degeneration of the PL seemsto follow a decreased tendon associated with modification inthe type of belly fibers from pennate to fusiform therefore inprimates with more derived thoracic limbs the weaker PL

To our knowledge this is the first study to present anevolutionary anatomical comparison of the PL Further stud-ies could indicate more accurately the evolutionary pathway

6 BioMed Research International

followed by PL from primitive to derived primates associatedwith aspects such as the muscular force difference betweenprimates and verification of anatomical aspects in morespecies and consider intra- and interspecific variations

5 Conclusions

Apparently in primates the PL tendon relative size seems toincrease from ancestors genera to more derived ones Thissuggests that this muscle is weaker in terrestrial primateswhen compared to arboreal primates The PL tendon seemsto be more derived in Gorilla sp considering the factorsanalyzed here but the tendon is longer in modern humans

Conflict of Interests

The authors declare that there is no conflict of interestsregarding the publication of this paper

References

[1] S Standring ldquoPelvis girdle and lower limbrdquo in Grayrsquos Anatomythe Anatomical Basis of Clinical Practice Churchill LivingstoneLondon UK 2008

[2] J W M Kigera and S Mukwaya ldquoFrequency of agenesisPalmaris longus through clinical examinationmdashan East Africanstudyrdquo PLoS ONE vol 6 no 12 Article ID e28997 2011

[3] H Gray ldquoMembro superiorrdquo in Gray Anatomia GuanabaraKoogan Rio de Janeiro Brazil 1979

[4] V S Telles and B A L Ernesto ldquoConsideraciones anatomicasde los musculos inconstantesrdquo MedUnab vol 1 no 3 pp 165ndash170 1998

[5] J M Cope E M Looney C A Craig R Gawron R Lamprosand R Mahoney ldquoMedian nerve compression and the reversedpalmaris longusrdquo International Journal od Anatomical Varia-tion vol 2 pp 102ndash104 2009

[6] A B M Machado and L J A Di Dio ldquoFrequency of themusculus palmaris longus studied in vivo in some Amazonindiansrdquo American Journal of Physical Anthropology vol 27 no1 pp 11ndash20 1967

[7] M A Babinski K M A Valente C M S Savedra C H FBurity and A M S Sabra ldquoVariacao anatomica do musculopalmar longo associado a ausencia do arco palmar superficiale compressao do canal de GuyonrdquoActa Scienteae Medica vol 2no 2 pp 54ndash58 2008

[8] C Fowlie C Fuller and M K Pratten ldquoAssessment of thepresenceabsence of the palmaris longus muscle in differentsports and elite andnon-elite sport populationsrdquoPhysiotherapyvol 98 no 2 pp 138ndash142 2012

[9] L Testut and A Latarjet Tratado de Anatomia Humana SalvatEditores Barcelona Espana 1959

[10] S Gibbs Comparative soft tissue morphology of the extantHominoidea including man [PhD thesis] The University ofLiverpool Inglaterra 1999

[11] A O Kayode A A Olamide I O Blessing and O U VictorldquoIncidence of palmaris longus muscle absence in Nigerianpopulationrdquo International Journal of Morphology vol 26 no 2pp 305ndash308 2008

[12] L C Angelini Junior F B Angelini B C Oliveira S ASoares L C Angelini and R H Cabral ldquoUse of the tendon

of the palmaris longus muscle in surgical proedures study oncadaversrdquoActaOrtopedica Brasileira vol 20 no 4 pp 226ndash2292012

[13] T A Aversi-Ferreira L G Vieira R M Pires Z Silvaand N Penha-Silva ldquoEstudo anatomico dos musculos flexoressuperficiais do antebraco no macaco Cebus apellardquo BioscienceJournal vol 22 no 1 pp 139ndash144 2006

[14] T A Aversi-Ferreira ldquoA new statistical method for comparativeanatomyrdquo International Journal of Morphology vol 27 no 4 pp1051ndash1058 2009

[15] T A Aversi-Ferreira R S Maior F O Carneiro-e-Silva et alldquoComparative anatomical analyses of the forearm muscles ofCebus libidinosus (Rylands et al 2000) manipulatory behaviorand tool userdquo PLoS ONE vol 6 no 7 Article ID e22165 8 pages2011

[16] H Preuschoft ldquoMuscles and joints of the anterior extremity ofa gorilla (Gorilla gorilla Savage et Wyman 1847)rdquo Gegenbaursmorphologisches Jahrbuch vol 107 no 2 pp 99ndash183 1965

[17] E E Sarmiento ldquoTerrestrial traits in the hands and feet ofgorillasrdquo American Museum Novitates vol 3091 pp 1ndash56

[18] R Diogo J F Pastor E M Ferrero et al Photographic andDescriptive Musculoskeletal Atlas of Gorilla With Notes on theAttachments Veriations Innervation Synonymy and Weight ofthe Muscles Science Publisher Enfield USA 2010

[19] A Keith ldquoOn the chimpanzees and their relationship to thegorillardquo Proceedings of the Zoological Society of London vol 67no 2 pp 296ndash312 1899

[20] E Loth Anthropologie Des Parties Molles (Muscles IntestinsVaisseaux Nerfs Peripheriques) Mianowski-Masson et CieParis France 1931

[21] G T Meglioli ldquoEtude de Irsquoheredite du muscle petit palmaire(Musculus palmaris longus) dans la population belgerdquo BullEtinde La Societe Royale Belge DrsquoAnthropologie et de Prehistoire vol75 pp 67ndash86 1965

[22] PW Brand R B Beach and D EThompson ldquoRelative tensionand potential excursion of muscles in the forearm and handrdquoJournal of Hand Surgery vol 6 no 3 pp 210ndash219 1981

[23] F Ankel-Simons Primate Anatomy An Introduction AcademicPress Orlando Fla USA 2000

[24] Y Ichinose H KanehisaM Ito Y Kawakami and T FukunagaldquoRelationship between muscle fiber pennation and force gen-eration capability in Olympic athletesrdquo International Journal ofSports Medicine vol 19 no 8 pp 541ndash546 1998

Submit your manuscripts athttpwwwhindawicom

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

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International Journal of

Volume 2014

Zoology

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Molecular Biology International

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The Scientific World JournalHindawi Publishing Corporation httpwwwhindawicom Volume 2014

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BioinformaticsAdvances in

Marine BiologyJournal of

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Signal TransductionJournal of

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BioMed Research International

Evolutionary BiologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Biochemistry Research International

ArchaeaHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

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Virolog y

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Nucleic AcidsJournal of

Volume 2014

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International Journal of

Microbiology

Page 3: Research Article Morphometric and Statistical Analysis of ...downloads.hindawi.com/journals/bmri/2014/178906.pdf · frequency or case studies, but comparative anatomy in primates

BioMed Research International 3

(a) (b) (c) (d) (e) (f)

(g) (h) (i)

Figure 1 Photos of the forearms of (a) Propithecus sp (045x left) (b) Lemur catta (052x left) (c) Sapajus libidinosus (034x right) (d)Atelessp (01x right) (e) Callithrix sp (03x right) (f) Aotus sp (08x right) (g) Macaca fuscata (034x right) (h) Pongo sp (079x left) (i) Pansp (023x left) From (a) to (f) muscles are pennate and from (g) to (i) are fusiform lowast indicates the palmaris longus

calculated or Comparative Anatomy Index (CAI) betweensamples from different species

CAI = 1003816100381610038161003816

PAF119894minus PAF

119894119894

1003816100381610038161003816

(3)

where indices 119894 and 119894119894 represent samples 1 and 2From the previous equations it follows that CAI value

close to zero represents greater similarity between sam-ples whereas CAI closer to 10 implies higher divergencebetween samples Therefore the greater the numerical dis-tance between values the greater the divergence in relationto the palmaris longus muscle between species

3 Results

In all primates studied here the PL tendon originated frommedial humeral epicondyle and inserted into the palmar

fascia was innervated by the median nerve A close relation-ship between the palmaris longus tendon and the fascia ofthe forearm was observed which is similar to other flexormuscles of the forearm The belly of palmaris longus waseasily distinguished from the other muscles of the forearmin all studied species except for the only exemplar of Atelessp in which the flexor muscles formed one group of belliesfrom the elbowwith a separation of tendons close to the wrist(Figure 1)

Regarding the type of fiber arrangement in the belly thepalmaris longus tendon presented an aspect similar to pen-nate in all Lemuriformes and NewWorld primates (Figure 1)but fusiform to Macaca fuscata (unique exemplar of OldWorld primates) and to apes (Pongo sp and Pan sp) andmodern humansThe average palmaris longustendon lengthshowed significant differences between species studied in thiswork (Table 1 Figure 2) Shorter tendons were observed in

4 BioMed Research International

Table 1 Measures of the palmaris longus palmaris longus tendon and palmaris longustendon relationship for species of primates andprimate groups

Specimens (119899) Total Tendon Totaltendon Primate groupsModern humans (28) 2551 (plusmn169) 1506 (plusmn175) 171 (plusmn013) Modern humansdaggerΔlowast+

Pan sp (3) 883 (plusmn162) 495 (plusmn091) 178 (plusmn004) ApesdaggerΔlowast+Pongo sp (2) 830 (plusmn010) 440 (plusmn030) 189 (plusmn015)Macaca fuscata (6) 1730 (plusmn043) 730 (plusmn05) 237 (plusmn012) Old World primatesdaggerΔlowast

Aotus sp (4) 68 (plusmn047) 32 (plusmn050) 209 (plusmn023)

NewWorld primatesdaggerΔCallithrix sp (3) 435 (plusmn019) 172 (plusmn023) 253 (plusmn008)Ateles sp (1) 253 100 253Sapajus libidinosus (11) 1071 (plusmn150) 299 (plusmn051) 381 (plusmn107)Lemur catta (2) 86 (plusmn050) 19 (plusmn00) 453 (plusmn027) LemuriformesdaggerPropithecus sp (2) 1125 (plusmn005) 22 (plusmn020) 516 (plusmn049)daggerSignificant difference among Lemuriformes and other groupsΔSignificant difference among NewWorld primates and other groupslowast Significant difference among Old World primates and other groups+Significant difference between apes and modern humans

Table 2 Statistics of the comparative anatomy of palmaris longus

Taxon PAF PAF =RF1 times 1198751 + RF2 times 1198752 + RF3 times 1198753 + RF4 times 1198754 + RF5 times 1198755

1198751+ 1198752+ 1198753+ 1198754+ 1198755

CAI = |PAF119894minus PAF

119894119894|

Propithecus sp 1 PAF = 4 times 1 + 4 times 1 + 3 times 1 + 2 times 1 + 1 times 114

Reference (most primitive characters)

Lemur catta 1 PAF = 4 times 1 + 4 times 1 + 3 times 1 + 2 times 1 + 1 times 114

0

Callithrix sp 1 PAF = 4 times 1 + 4 times 1 + 3 times 1 + 2 times 1 + 1 times 114

0

Sapajus sp 1 PAF = 4 times 1 + 4 times 1 + 3 times 1 + 2 times 1 + 1 times 114

0

Ateles sp 1 PAF = 4 times 1 + 4 times 1 + 3 times 1 + 2 times 1 + 1 times 114

0

Macaca fuscata 0964 PAF = 4 times 1 + 4 times 1 + 3 times 1 + 2 times 1 + 1 times 0514

0036

Pongo sp 0964 PAF = 4 times 1 + 4 times 1 + 3 times 1 + 2 times 1 + 1 times 0514

0036

Modern humans 0947 PAF = 4 times 1 + 4 times 1 + 3 times 1 + 2 times 0878 + 1 times 0514

0053

Pan sp 0918 PAF = 4 times 1 + 4 times 1 + 3 times 1 + 2 times 0678 + 1 times 0514

0082

Gorilla sp 0867 PAF = 4 times 1 + 4 times 1 + 3 times 1 + 2 times 0316 + 1 times 0514

0133

PAF is the weighted average of frequencies CAI is Comparative Anatomy Index

Lemuriformes and longer in modern humans (119875 lt 005Figure 2)

Significant differences were observed between averages(119875 lt 005) calculated for all groups that is Lemuriformes andother groups NewWorld primates and other groups andOldWorld primates (Macaca fuscata) and other groups and forapes andmodern humans (Table 1 Figure 2) Nevertheless toAotus sp specifically in comparison to Macaca fuscata andapes differences were not observed

The CAI indicates that Lemuriformes and New Worldprimates share similar characters to PL (CAI = 0) while

these features are more derived in Gorilla sp (CAI = 0133)(Table 2)

4 Discussion

The large variations in the prevalence of the PL amongmodern humans may be indicative that this muscle is degen-erating [1] and its small belly may suggest it is a vestigialmuscle [2]

Although there are several studies investigating the fre-quency of PL in modern humans only a few recent studies

BioMed Research International 5

Lemuriformes New Worldprimates primates

Old World Apes Modernhumans

BoxMean line

0

1

2

3

4

5

6

Palm

aris

long

uste

ndon

(cm

)

Figure 2 Graph showing the mean line relative to the palmarislongustendon length of primate groups

have compared PL frequency in nonhuman primates [9 1016ndash18] From such studies it was found for example that thefrequency of PL in Gorilla sp is much lower than in humansand chimpanzees even though humans and chimpanzees arethe only species in which absence of PL has been reportedThe PL is found in Gorilla sp from 15 to 63 accordingto some authors [10 16 17 19 20] and could be consideredabsent in gorillas according to [18]

In this sense considering only the frequency of thismuscle in primates PLwould bemore degenerative inGorillasp Also if degeneration is considered a derived characteristicof PL it is more derived in Gorilla sp than in other primatesand modern humans Indeed it is reasonable to suggest thatPL is a degenerative muscle because according to Meglioli[21] its agenesis is associated with a recessive gene

Notwithstanding when parameters such as origin inser-tion innervation frequency and type of belly fibers arecompared together and using specific weight (ie whenCAI is applied for review see Aversi-Ferreira [14] andAversi-Ferreira et al [15]) Gorilla sp shows the greatestnumerical distance from the primitive character consideredin Propithecus sp followed by Pan sp modern humansPongo sp and Macaca fuscata in decreasing order

In fact the different variation factor observed in the CAIcalculation can be explained by the low frequency in thenumber of specimens and species and by the different types ofmuscle fibers of the belly muscle obtained from the literatureand from the present study To Lemuriformes andNewWorldprimates CAI indicates no quantitative difference that is thecalculated value is zero but toMacaca fuscata and Pongo spthe only difference is type of fiber arrangement

In order to obtain more objective parameters the relativelength of the tendon was studied that is the length of thePL muscle divided by the length of the palmaris longus

tendon Its measure indicates indirectly the muscle relativestrength because smaller belly indicates less sarcomeresacting together to generate contraction [22] Therefore alonger tendon indicates less muscular force According tothe data obtained here this relation (PL muscle divided bythe length of the palmaris longus tendon) decreases fromLemuriformes to modern humans

Interestingly the comparison between means of therelative measures of PL among these groups of primates(119875 lt 005) showed significant differences between one groupand all other groups One discrepancy was observed in themeasures obtained from Aotus sp a New World primate inwhich the average measures were similar to Macaca fuscataand apes but it was not possible to associate it with any aspectconsidered in this work Putatively a weak PL is observedin modern humans and apes and strong in LemuriformesTherefore it is possible to hypothesize that the tendon lengthdecreases from arboreal to terrestrial primates In line withthis Ankel-simons [23] reports that great apes move in aquadrupedal Knuckle-walking manner the Macaca genusis terrestrial quadruped all New World primates are highlyarboreal and all Lemuriformes except for Lemur catta spendaround 13 of the time on the ground

Specifically to modern humans the average measure ofthe PL tendon obtained here that is 1506 (plusmn175) wasdifferent from those reported for other Brazilians cadavers[12] (1199 plusmn 152) Apparently these differences are due tocriteria for length measurements of the palmaris longustendon in both studies

On the other hand different fiber arrangement of thebelly of PL in Lemuriformes and New World primatescompared to OldWorld primates apes and modern humanswas observed The tendon begins in the belly which involvesthe tendon laterally up to approximately the wrist in Lemu-riformes and New World primates characterizing a pennatemuscle whereas fromMacaca fuscata tomodern humans thetendon begins after the end of the belly observed via nakedeye characterizing a fusiform muscle

This characteristic affects the muscular strength thefusiform muscle generates a more direct contraction butto modern humans apes and Macaca fuscata the belliesare shorter in a pennate muscle however the physiologicalcross-sectional area is considerably larger than its anatomicalcross-sectional area Therefore the pennate muscle ceterisparibus generates more force [24] In addition they arelonger in Lemuriformes and New World primate than inother primates

The number of animals per species and the number ofspecies used here allowed inferring the following conclusionsFirst the more derived characteristics of PL are found inapes especially in Gorilla sp relative to evolutionary aspectsmeasured by CAI but a shorter tendon is verified in modernhumans Second the path to the degeneration of the PL seemsto follow a decreased tendon associated with modification inthe type of belly fibers from pennate to fusiform therefore inprimates with more derived thoracic limbs the weaker PL

To our knowledge this is the first study to present anevolutionary anatomical comparison of the PL Further stud-ies could indicate more accurately the evolutionary pathway

6 BioMed Research International

followed by PL from primitive to derived primates associatedwith aspects such as the muscular force difference betweenprimates and verification of anatomical aspects in morespecies and consider intra- and interspecific variations

5 Conclusions

Apparently in primates the PL tendon relative size seems toincrease from ancestors genera to more derived ones Thissuggests that this muscle is weaker in terrestrial primateswhen compared to arboreal primates The PL tendon seemsto be more derived in Gorilla sp considering the factorsanalyzed here but the tendon is longer in modern humans

Conflict of Interests

The authors declare that there is no conflict of interestsregarding the publication of this paper

References

[1] S Standring ldquoPelvis girdle and lower limbrdquo in Grayrsquos Anatomythe Anatomical Basis of Clinical Practice Churchill LivingstoneLondon UK 2008

[2] J W M Kigera and S Mukwaya ldquoFrequency of agenesisPalmaris longus through clinical examinationmdashan East Africanstudyrdquo PLoS ONE vol 6 no 12 Article ID e28997 2011

[3] H Gray ldquoMembro superiorrdquo in Gray Anatomia GuanabaraKoogan Rio de Janeiro Brazil 1979

[4] V S Telles and B A L Ernesto ldquoConsideraciones anatomicasde los musculos inconstantesrdquo MedUnab vol 1 no 3 pp 165ndash170 1998

[5] J M Cope E M Looney C A Craig R Gawron R Lamprosand R Mahoney ldquoMedian nerve compression and the reversedpalmaris longusrdquo International Journal od Anatomical Varia-tion vol 2 pp 102ndash104 2009

[6] A B M Machado and L J A Di Dio ldquoFrequency of themusculus palmaris longus studied in vivo in some Amazonindiansrdquo American Journal of Physical Anthropology vol 27 no1 pp 11ndash20 1967

[7] M A Babinski K M A Valente C M S Savedra C H FBurity and A M S Sabra ldquoVariacao anatomica do musculopalmar longo associado a ausencia do arco palmar superficiale compressao do canal de GuyonrdquoActa Scienteae Medica vol 2no 2 pp 54ndash58 2008

[8] C Fowlie C Fuller and M K Pratten ldquoAssessment of thepresenceabsence of the palmaris longus muscle in differentsports and elite andnon-elite sport populationsrdquoPhysiotherapyvol 98 no 2 pp 138ndash142 2012

[9] L Testut and A Latarjet Tratado de Anatomia Humana SalvatEditores Barcelona Espana 1959

[10] S Gibbs Comparative soft tissue morphology of the extantHominoidea including man [PhD thesis] The University ofLiverpool Inglaterra 1999

[11] A O Kayode A A Olamide I O Blessing and O U VictorldquoIncidence of palmaris longus muscle absence in Nigerianpopulationrdquo International Journal of Morphology vol 26 no 2pp 305ndash308 2008

[12] L C Angelini Junior F B Angelini B C Oliveira S ASoares L C Angelini and R H Cabral ldquoUse of the tendon

of the palmaris longus muscle in surgical proedures study oncadaversrdquoActaOrtopedica Brasileira vol 20 no 4 pp 226ndash2292012

[13] T A Aversi-Ferreira L G Vieira R M Pires Z Silvaand N Penha-Silva ldquoEstudo anatomico dos musculos flexoressuperficiais do antebraco no macaco Cebus apellardquo BioscienceJournal vol 22 no 1 pp 139ndash144 2006

[14] T A Aversi-Ferreira ldquoA new statistical method for comparativeanatomyrdquo International Journal of Morphology vol 27 no 4 pp1051ndash1058 2009

[15] T A Aversi-Ferreira R S Maior F O Carneiro-e-Silva et alldquoComparative anatomical analyses of the forearm muscles ofCebus libidinosus (Rylands et al 2000) manipulatory behaviorand tool userdquo PLoS ONE vol 6 no 7 Article ID e22165 8 pages2011

[16] H Preuschoft ldquoMuscles and joints of the anterior extremity ofa gorilla (Gorilla gorilla Savage et Wyman 1847)rdquo Gegenbaursmorphologisches Jahrbuch vol 107 no 2 pp 99ndash183 1965

[17] E E Sarmiento ldquoTerrestrial traits in the hands and feet ofgorillasrdquo American Museum Novitates vol 3091 pp 1ndash56

[18] R Diogo J F Pastor E M Ferrero et al Photographic andDescriptive Musculoskeletal Atlas of Gorilla With Notes on theAttachments Veriations Innervation Synonymy and Weight ofthe Muscles Science Publisher Enfield USA 2010

[19] A Keith ldquoOn the chimpanzees and their relationship to thegorillardquo Proceedings of the Zoological Society of London vol 67no 2 pp 296ndash312 1899

[20] E Loth Anthropologie Des Parties Molles (Muscles IntestinsVaisseaux Nerfs Peripheriques) Mianowski-Masson et CieParis France 1931

[21] G T Meglioli ldquoEtude de Irsquoheredite du muscle petit palmaire(Musculus palmaris longus) dans la population belgerdquo BullEtinde La Societe Royale Belge DrsquoAnthropologie et de Prehistoire vol75 pp 67ndash86 1965

[22] PW Brand R B Beach and D EThompson ldquoRelative tensionand potential excursion of muscles in the forearm and handrdquoJournal of Hand Surgery vol 6 no 3 pp 210ndash219 1981

[23] F Ankel-Simons Primate Anatomy An Introduction AcademicPress Orlando Fla USA 2000

[24] Y Ichinose H KanehisaM Ito Y Kawakami and T FukunagaldquoRelationship between muscle fiber pennation and force gen-eration capability in Olympic athletesrdquo International Journal ofSports Medicine vol 19 no 8 pp 541ndash546 1998

Submit your manuscripts athttpwwwhindawicom

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Anatomy Research International

PeptidesInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporation httpwwwhindawicom

International Journal of

Volume 2014

Zoology

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Molecular Biology International

GenomicsInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

The Scientific World JournalHindawi Publishing Corporation httpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BioinformaticsAdvances in

Marine BiologyJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Signal TransductionJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BioMed Research International

Evolutionary BiologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Biochemistry Research International

ArchaeaHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Genetics Research International

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Advances in

Virolog y

Hindawi Publishing Corporationhttpwwwhindawicom

Nucleic AcidsJournal of

Volume 2014

Stem CellsInternational

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Enzyme Research

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

International Journal of

Microbiology

Page 4: Research Article Morphometric and Statistical Analysis of ...downloads.hindawi.com/journals/bmri/2014/178906.pdf · frequency or case studies, but comparative anatomy in primates

4 BioMed Research International

Table 1 Measures of the palmaris longus palmaris longus tendon and palmaris longustendon relationship for species of primates andprimate groups

Specimens (119899) Total Tendon Totaltendon Primate groupsModern humans (28) 2551 (plusmn169) 1506 (plusmn175) 171 (plusmn013) Modern humansdaggerΔlowast+

Pan sp (3) 883 (plusmn162) 495 (plusmn091) 178 (plusmn004) ApesdaggerΔlowast+Pongo sp (2) 830 (plusmn010) 440 (plusmn030) 189 (plusmn015)Macaca fuscata (6) 1730 (plusmn043) 730 (plusmn05) 237 (plusmn012) Old World primatesdaggerΔlowast

Aotus sp (4) 68 (plusmn047) 32 (plusmn050) 209 (plusmn023)

NewWorld primatesdaggerΔCallithrix sp (3) 435 (plusmn019) 172 (plusmn023) 253 (plusmn008)Ateles sp (1) 253 100 253Sapajus libidinosus (11) 1071 (plusmn150) 299 (plusmn051) 381 (plusmn107)Lemur catta (2) 86 (plusmn050) 19 (plusmn00) 453 (plusmn027) LemuriformesdaggerPropithecus sp (2) 1125 (plusmn005) 22 (plusmn020) 516 (plusmn049)daggerSignificant difference among Lemuriformes and other groupsΔSignificant difference among NewWorld primates and other groupslowast Significant difference among Old World primates and other groups+Significant difference between apes and modern humans

Table 2 Statistics of the comparative anatomy of palmaris longus

Taxon PAF PAF =RF1 times 1198751 + RF2 times 1198752 + RF3 times 1198753 + RF4 times 1198754 + RF5 times 1198755

1198751+ 1198752+ 1198753+ 1198754+ 1198755

CAI = |PAF119894minus PAF

119894119894|

Propithecus sp 1 PAF = 4 times 1 + 4 times 1 + 3 times 1 + 2 times 1 + 1 times 114

Reference (most primitive characters)

Lemur catta 1 PAF = 4 times 1 + 4 times 1 + 3 times 1 + 2 times 1 + 1 times 114

0

Callithrix sp 1 PAF = 4 times 1 + 4 times 1 + 3 times 1 + 2 times 1 + 1 times 114

0

Sapajus sp 1 PAF = 4 times 1 + 4 times 1 + 3 times 1 + 2 times 1 + 1 times 114

0

Ateles sp 1 PAF = 4 times 1 + 4 times 1 + 3 times 1 + 2 times 1 + 1 times 114

0

Macaca fuscata 0964 PAF = 4 times 1 + 4 times 1 + 3 times 1 + 2 times 1 + 1 times 0514

0036

Pongo sp 0964 PAF = 4 times 1 + 4 times 1 + 3 times 1 + 2 times 1 + 1 times 0514

0036

Modern humans 0947 PAF = 4 times 1 + 4 times 1 + 3 times 1 + 2 times 0878 + 1 times 0514

0053

Pan sp 0918 PAF = 4 times 1 + 4 times 1 + 3 times 1 + 2 times 0678 + 1 times 0514

0082

Gorilla sp 0867 PAF = 4 times 1 + 4 times 1 + 3 times 1 + 2 times 0316 + 1 times 0514

0133

PAF is the weighted average of frequencies CAI is Comparative Anatomy Index

Lemuriformes and longer in modern humans (119875 lt 005Figure 2)

Significant differences were observed between averages(119875 lt 005) calculated for all groups that is Lemuriformes andother groups NewWorld primates and other groups andOldWorld primates (Macaca fuscata) and other groups and forapes andmodern humans (Table 1 Figure 2) Nevertheless toAotus sp specifically in comparison to Macaca fuscata andapes differences were not observed

The CAI indicates that Lemuriformes and New Worldprimates share similar characters to PL (CAI = 0) while

these features are more derived in Gorilla sp (CAI = 0133)(Table 2)

4 Discussion

The large variations in the prevalence of the PL amongmodern humans may be indicative that this muscle is degen-erating [1] and its small belly may suggest it is a vestigialmuscle [2]

Although there are several studies investigating the fre-quency of PL in modern humans only a few recent studies

BioMed Research International 5

Lemuriformes New Worldprimates primates

Old World Apes Modernhumans

BoxMean line

0

1

2

3

4

5

6

Palm

aris

long

uste

ndon

(cm

)

Figure 2 Graph showing the mean line relative to the palmarislongustendon length of primate groups

have compared PL frequency in nonhuman primates [9 1016ndash18] From such studies it was found for example that thefrequency of PL in Gorilla sp is much lower than in humansand chimpanzees even though humans and chimpanzees arethe only species in which absence of PL has been reportedThe PL is found in Gorilla sp from 15 to 63 accordingto some authors [10 16 17 19 20] and could be consideredabsent in gorillas according to [18]

In this sense considering only the frequency of thismuscle in primates PLwould bemore degenerative inGorillasp Also if degeneration is considered a derived characteristicof PL it is more derived in Gorilla sp than in other primatesand modern humans Indeed it is reasonable to suggest thatPL is a degenerative muscle because according to Meglioli[21] its agenesis is associated with a recessive gene

Notwithstanding when parameters such as origin inser-tion innervation frequency and type of belly fibers arecompared together and using specific weight (ie whenCAI is applied for review see Aversi-Ferreira [14] andAversi-Ferreira et al [15]) Gorilla sp shows the greatestnumerical distance from the primitive character consideredin Propithecus sp followed by Pan sp modern humansPongo sp and Macaca fuscata in decreasing order

In fact the different variation factor observed in the CAIcalculation can be explained by the low frequency in thenumber of specimens and species and by the different types ofmuscle fibers of the belly muscle obtained from the literatureand from the present study To Lemuriformes andNewWorldprimates CAI indicates no quantitative difference that is thecalculated value is zero but toMacaca fuscata and Pongo spthe only difference is type of fiber arrangement

In order to obtain more objective parameters the relativelength of the tendon was studied that is the length of thePL muscle divided by the length of the palmaris longus

tendon Its measure indicates indirectly the muscle relativestrength because smaller belly indicates less sarcomeresacting together to generate contraction [22] Therefore alonger tendon indicates less muscular force According tothe data obtained here this relation (PL muscle divided bythe length of the palmaris longus tendon) decreases fromLemuriformes to modern humans

Interestingly the comparison between means of therelative measures of PL among these groups of primates(119875 lt 005) showed significant differences between one groupand all other groups One discrepancy was observed in themeasures obtained from Aotus sp a New World primate inwhich the average measures were similar to Macaca fuscataand apes but it was not possible to associate it with any aspectconsidered in this work Putatively a weak PL is observedin modern humans and apes and strong in LemuriformesTherefore it is possible to hypothesize that the tendon lengthdecreases from arboreal to terrestrial primates In line withthis Ankel-simons [23] reports that great apes move in aquadrupedal Knuckle-walking manner the Macaca genusis terrestrial quadruped all New World primates are highlyarboreal and all Lemuriformes except for Lemur catta spendaround 13 of the time on the ground

Specifically to modern humans the average measure ofthe PL tendon obtained here that is 1506 (plusmn175) wasdifferent from those reported for other Brazilians cadavers[12] (1199 plusmn 152) Apparently these differences are due tocriteria for length measurements of the palmaris longustendon in both studies

On the other hand different fiber arrangement of thebelly of PL in Lemuriformes and New World primatescompared to OldWorld primates apes and modern humanswas observed The tendon begins in the belly which involvesthe tendon laterally up to approximately the wrist in Lemu-riformes and New World primates characterizing a pennatemuscle whereas fromMacaca fuscata tomodern humans thetendon begins after the end of the belly observed via nakedeye characterizing a fusiform muscle

This characteristic affects the muscular strength thefusiform muscle generates a more direct contraction butto modern humans apes and Macaca fuscata the belliesare shorter in a pennate muscle however the physiologicalcross-sectional area is considerably larger than its anatomicalcross-sectional area Therefore the pennate muscle ceterisparibus generates more force [24] In addition they arelonger in Lemuriformes and New World primate than inother primates

The number of animals per species and the number ofspecies used here allowed inferring the following conclusionsFirst the more derived characteristics of PL are found inapes especially in Gorilla sp relative to evolutionary aspectsmeasured by CAI but a shorter tendon is verified in modernhumans Second the path to the degeneration of the PL seemsto follow a decreased tendon associated with modification inthe type of belly fibers from pennate to fusiform therefore inprimates with more derived thoracic limbs the weaker PL

To our knowledge this is the first study to present anevolutionary anatomical comparison of the PL Further stud-ies could indicate more accurately the evolutionary pathway

6 BioMed Research International

followed by PL from primitive to derived primates associatedwith aspects such as the muscular force difference betweenprimates and verification of anatomical aspects in morespecies and consider intra- and interspecific variations

5 Conclusions

Apparently in primates the PL tendon relative size seems toincrease from ancestors genera to more derived ones Thissuggests that this muscle is weaker in terrestrial primateswhen compared to arboreal primates The PL tendon seemsto be more derived in Gorilla sp considering the factorsanalyzed here but the tendon is longer in modern humans

Conflict of Interests

The authors declare that there is no conflict of interestsregarding the publication of this paper

References

[1] S Standring ldquoPelvis girdle and lower limbrdquo in Grayrsquos Anatomythe Anatomical Basis of Clinical Practice Churchill LivingstoneLondon UK 2008

[2] J W M Kigera and S Mukwaya ldquoFrequency of agenesisPalmaris longus through clinical examinationmdashan East Africanstudyrdquo PLoS ONE vol 6 no 12 Article ID e28997 2011

[3] H Gray ldquoMembro superiorrdquo in Gray Anatomia GuanabaraKoogan Rio de Janeiro Brazil 1979

[4] V S Telles and B A L Ernesto ldquoConsideraciones anatomicasde los musculos inconstantesrdquo MedUnab vol 1 no 3 pp 165ndash170 1998

[5] J M Cope E M Looney C A Craig R Gawron R Lamprosand R Mahoney ldquoMedian nerve compression and the reversedpalmaris longusrdquo International Journal od Anatomical Varia-tion vol 2 pp 102ndash104 2009

[6] A B M Machado and L J A Di Dio ldquoFrequency of themusculus palmaris longus studied in vivo in some Amazonindiansrdquo American Journal of Physical Anthropology vol 27 no1 pp 11ndash20 1967

[7] M A Babinski K M A Valente C M S Savedra C H FBurity and A M S Sabra ldquoVariacao anatomica do musculopalmar longo associado a ausencia do arco palmar superficiale compressao do canal de GuyonrdquoActa Scienteae Medica vol 2no 2 pp 54ndash58 2008

[8] C Fowlie C Fuller and M K Pratten ldquoAssessment of thepresenceabsence of the palmaris longus muscle in differentsports and elite andnon-elite sport populationsrdquoPhysiotherapyvol 98 no 2 pp 138ndash142 2012

[9] L Testut and A Latarjet Tratado de Anatomia Humana SalvatEditores Barcelona Espana 1959

[10] S Gibbs Comparative soft tissue morphology of the extantHominoidea including man [PhD thesis] The University ofLiverpool Inglaterra 1999

[11] A O Kayode A A Olamide I O Blessing and O U VictorldquoIncidence of palmaris longus muscle absence in Nigerianpopulationrdquo International Journal of Morphology vol 26 no 2pp 305ndash308 2008

[12] L C Angelini Junior F B Angelini B C Oliveira S ASoares L C Angelini and R H Cabral ldquoUse of the tendon

of the palmaris longus muscle in surgical proedures study oncadaversrdquoActaOrtopedica Brasileira vol 20 no 4 pp 226ndash2292012

[13] T A Aversi-Ferreira L G Vieira R M Pires Z Silvaand N Penha-Silva ldquoEstudo anatomico dos musculos flexoressuperficiais do antebraco no macaco Cebus apellardquo BioscienceJournal vol 22 no 1 pp 139ndash144 2006

[14] T A Aversi-Ferreira ldquoA new statistical method for comparativeanatomyrdquo International Journal of Morphology vol 27 no 4 pp1051ndash1058 2009

[15] T A Aversi-Ferreira R S Maior F O Carneiro-e-Silva et alldquoComparative anatomical analyses of the forearm muscles ofCebus libidinosus (Rylands et al 2000) manipulatory behaviorand tool userdquo PLoS ONE vol 6 no 7 Article ID e22165 8 pages2011

[16] H Preuschoft ldquoMuscles and joints of the anterior extremity ofa gorilla (Gorilla gorilla Savage et Wyman 1847)rdquo Gegenbaursmorphologisches Jahrbuch vol 107 no 2 pp 99ndash183 1965

[17] E E Sarmiento ldquoTerrestrial traits in the hands and feet ofgorillasrdquo American Museum Novitates vol 3091 pp 1ndash56

[18] R Diogo J F Pastor E M Ferrero et al Photographic andDescriptive Musculoskeletal Atlas of Gorilla With Notes on theAttachments Veriations Innervation Synonymy and Weight ofthe Muscles Science Publisher Enfield USA 2010

[19] A Keith ldquoOn the chimpanzees and their relationship to thegorillardquo Proceedings of the Zoological Society of London vol 67no 2 pp 296ndash312 1899

[20] E Loth Anthropologie Des Parties Molles (Muscles IntestinsVaisseaux Nerfs Peripheriques) Mianowski-Masson et CieParis France 1931

[21] G T Meglioli ldquoEtude de Irsquoheredite du muscle petit palmaire(Musculus palmaris longus) dans la population belgerdquo BullEtinde La Societe Royale Belge DrsquoAnthropologie et de Prehistoire vol75 pp 67ndash86 1965

[22] PW Brand R B Beach and D EThompson ldquoRelative tensionand potential excursion of muscles in the forearm and handrdquoJournal of Hand Surgery vol 6 no 3 pp 210ndash219 1981

[23] F Ankel-Simons Primate Anatomy An Introduction AcademicPress Orlando Fla USA 2000

[24] Y Ichinose H KanehisaM Ito Y Kawakami and T FukunagaldquoRelationship between muscle fiber pennation and force gen-eration capability in Olympic athletesrdquo International Journal ofSports Medicine vol 19 no 8 pp 541ndash546 1998

Submit your manuscripts athttpwwwhindawicom

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Anatomy Research International

PeptidesInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporation httpwwwhindawicom

International Journal of

Volume 2014

Zoology

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Molecular Biology International

GenomicsInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

The Scientific World JournalHindawi Publishing Corporation httpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BioinformaticsAdvances in

Marine BiologyJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Signal TransductionJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BioMed Research International

Evolutionary BiologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Biochemistry Research International

ArchaeaHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Genetics Research International

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Advances in

Virolog y

Hindawi Publishing Corporationhttpwwwhindawicom

Nucleic AcidsJournal of

Volume 2014

Stem CellsInternational

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Enzyme Research

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

International Journal of

Microbiology

Page 5: Research Article Morphometric and Statistical Analysis of ...downloads.hindawi.com/journals/bmri/2014/178906.pdf · frequency or case studies, but comparative anatomy in primates

BioMed Research International 5

Lemuriformes New Worldprimates primates

Old World Apes Modernhumans

BoxMean line

0

1

2

3

4

5

6

Palm

aris

long

uste

ndon

(cm

)

Figure 2 Graph showing the mean line relative to the palmarislongustendon length of primate groups

have compared PL frequency in nonhuman primates [9 1016ndash18] From such studies it was found for example that thefrequency of PL in Gorilla sp is much lower than in humansand chimpanzees even though humans and chimpanzees arethe only species in which absence of PL has been reportedThe PL is found in Gorilla sp from 15 to 63 accordingto some authors [10 16 17 19 20] and could be consideredabsent in gorillas according to [18]

In this sense considering only the frequency of thismuscle in primates PLwould bemore degenerative inGorillasp Also if degeneration is considered a derived characteristicof PL it is more derived in Gorilla sp than in other primatesand modern humans Indeed it is reasonable to suggest thatPL is a degenerative muscle because according to Meglioli[21] its agenesis is associated with a recessive gene

Notwithstanding when parameters such as origin inser-tion innervation frequency and type of belly fibers arecompared together and using specific weight (ie whenCAI is applied for review see Aversi-Ferreira [14] andAversi-Ferreira et al [15]) Gorilla sp shows the greatestnumerical distance from the primitive character consideredin Propithecus sp followed by Pan sp modern humansPongo sp and Macaca fuscata in decreasing order

In fact the different variation factor observed in the CAIcalculation can be explained by the low frequency in thenumber of specimens and species and by the different types ofmuscle fibers of the belly muscle obtained from the literatureand from the present study To Lemuriformes andNewWorldprimates CAI indicates no quantitative difference that is thecalculated value is zero but toMacaca fuscata and Pongo spthe only difference is type of fiber arrangement

In order to obtain more objective parameters the relativelength of the tendon was studied that is the length of thePL muscle divided by the length of the palmaris longus

tendon Its measure indicates indirectly the muscle relativestrength because smaller belly indicates less sarcomeresacting together to generate contraction [22] Therefore alonger tendon indicates less muscular force According tothe data obtained here this relation (PL muscle divided bythe length of the palmaris longus tendon) decreases fromLemuriformes to modern humans

Interestingly the comparison between means of therelative measures of PL among these groups of primates(119875 lt 005) showed significant differences between one groupand all other groups One discrepancy was observed in themeasures obtained from Aotus sp a New World primate inwhich the average measures were similar to Macaca fuscataand apes but it was not possible to associate it with any aspectconsidered in this work Putatively a weak PL is observedin modern humans and apes and strong in LemuriformesTherefore it is possible to hypothesize that the tendon lengthdecreases from arboreal to terrestrial primates In line withthis Ankel-simons [23] reports that great apes move in aquadrupedal Knuckle-walking manner the Macaca genusis terrestrial quadruped all New World primates are highlyarboreal and all Lemuriformes except for Lemur catta spendaround 13 of the time on the ground

Specifically to modern humans the average measure ofthe PL tendon obtained here that is 1506 (plusmn175) wasdifferent from those reported for other Brazilians cadavers[12] (1199 plusmn 152) Apparently these differences are due tocriteria for length measurements of the palmaris longustendon in both studies

On the other hand different fiber arrangement of thebelly of PL in Lemuriformes and New World primatescompared to OldWorld primates apes and modern humanswas observed The tendon begins in the belly which involvesthe tendon laterally up to approximately the wrist in Lemu-riformes and New World primates characterizing a pennatemuscle whereas fromMacaca fuscata tomodern humans thetendon begins after the end of the belly observed via nakedeye characterizing a fusiform muscle

This characteristic affects the muscular strength thefusiform muscle generates a more direct contraction butto modern humans apes and Macaca fuscata the belliesare shorter in a pennate muscle however the physiologicalcross-sectional area is considerably larger than its anatomicalcross-sectional area Therefore the pennate muscle ceterisparibus generates more force [24] In addition they arelonger in Lemuriformes and New World primate than inother primates

The number of animals per species and the number ofspecies used here allowed inferring the following conclusionsFirst the more derived characteristics of PL are found inapes especially in Gorilla sp relative to evolutionary aspectsmeasured by CAI but a shorter tendon is verified in modernhumans Second the path to the degeneration of the PL seemsto follow a decreased tendon associated with modification inthe type of belly fibers from pennate to fusiform therefore inprimates with more derived thoracic limbs the weaker PL

To our knowledge this is the first study to present anevolutionary anatomical comparison of the PL Further stud-ies could indicate more accurately the evolutionary pathway

6 BioMed Research International

followed by PL from primitive to derived primates associatedwith aspects such as the muscular force difference betweenprimates and verification of anatomical aspects in morespecies and consider intra- and interspecific variations

5 Conclusions

Apparently in primates the PL tendon relative size seems toincrease from ancestors genera to more derived ones Thissuggests that this muscle is weaker in terrestrial primateswhen compared to arboreal primates The PL tendon seemsto be more derived in Gorilla sp considering the factorsanalyzed here but the tendon is longer in modern humans

Conflict of Interests

The authors declare that there is no conflict of interestsregarding the publication of this paper

References

[1] S Standring ldquoPelvis girdle and lower limbrdquo in Grayrsquos Anatomythe Anatomical Basis of Clinical Practice Churchill LivingstoneLondon UK 2008

[2] J W M Kigera and S Mukwaya ldquoFrequency of agenesisPalmaris longus through clinical examinationmdashan East Africanstudyrdquo PLoS ONE vol 6 no 12 Article ID e28997 2011

[3] H Gray ldquoMembro superiorrdquo in Gray Anatomia GuanabaraKoogan Rio de Janeiro Brazil 1979

[4] V S Telles and B A L Ernesto ldquoConsideraciones anatomicasde los musculos inconstantesrdquo MedUnab vol 1 no 3 pp 165ndash170 1998

[5] J M Cope E M Looney C A Craig R Gawron R Lamprosand R Mahoney ldquoMedian nerve compression and the reversedpalmaris longusrdquo International Journal od Anatomical Varia-tion vol 2 pp 102ndash104 2009

[6] A B M Machado and L J A Di Dio ldquoFrequency of themusculus palmaris longus studied in vivo in some Amazonindiansrdquo American Journal of Physical Anthropology vol 27 no1 pp 11ndash20 1967

[7] M A Babinski K M A Valente C M S Savedra C H FBurity and A M S Sabra ldquoVariacao anatomica do musculopalmar longo associado a ausencia do arco palmar superficiale compressao do canal de GuyonrdquoActa Scienteae Medica vol 2no 2 pp 54ndash58 2008

[8] C Fowlie C Fuller and M K Pratten ldquoAssessment of thepresenceabsence of the palmaris longus muscle in differentsports and elite andnon-elite sport populationsrdquoPhysiotherapyvol 98 no 2 pp 138ndash142 2012

[9] L Testut and A Latarjet Tratado de Anatomia Humana SalvatEditores Barcelona Espana 1959

[10] S Gibbs Comparative soft tissue morphology of the extantHominoidea including man [PhD thesis] The University ofLiverpool Inglaterra 1999

[11] A O Kayode A A Olamide I O Blessing and O U VictorldquoIncidence of palmaris longus muscle absence in Nigerianpopulationrdquo International Journal of Morphology vol 26 no 2pp 305ndash308 2008

[12] L C Angelini Junior F B Angelini B C Oliveira S ASoares L C Angelini and R H Cabral ldquoUse of the tendon

of the palmaris longus muscle in surgical proedures study oncadaversrdquoActaOrtopedica Brasileira vol 20 no 4 pp 226ndash2292012

[13] T A Aversi-Ferreira L G Vieira R M Pires Z Silvaand N Penha-Silva ldquoEstudo anatomico dos musculos flexoressuperficiais do antebraco no macaco Cebus apellardquo BioscienceJournal vol 22 no 1 pp 139ndash144 2006

[14] T A Aversi-Ferreira ldquoA new statistical method for comparativeanatomyrdquo International Journal of Morphology vol 27 no 4 pp1051ndash1058 2009

[15] T A Aversi-Ferreira R S Maior F O Carneiro-e-Silva et alldquoComparative anatomical analyses of the forearm muscles ofCebus libidinosus (Rylands et al 2000) manipulatory behaviorand tool userdquo PLoS ONE vol 6 no 7 Article ID e22165 8 pages2011

[16] H Preuschoft ldquoMuscles and joints of the anterior extremity ofa gorilla (Gorilla gorilla Savage et Wyman 1847)rdquo Gegenbaursmorphologisches Jahrbuch vol 107 no 2 pp 99ndash183 1965

[17] E E Sarmiento ldquoTerrestrial traits in the hands and feet ofgorillasrdquo American Museum Novitates vol 3091 pp 1ndash56

[18] R Diogo J F Pastor E M Ferrero et al Photographic andDescriptive Musculoskeletal Atlas of Gorilla With Notes on theAttachments Veriations Innervation Synonymy and Weight ofthe Muscles Science Publisher Enfield USA 2010

[19] A Keith ldquoOn the chimpanzees and their relationship to thegorillardquo Proceedings of the Zoological Society of London vol 67no 2 pp 296ndash312 1899

[20] E Loth Anthropologie Des Parties Molles (Muscles IntestinsVaisseaux Nerfs Peripheriques) Mianowski-Masson et CieParis France 1931

[21] G T Meglioli ldquoEtude de Irsquoheredite du muscle petit palmaire(Musculus palmaris longus) dans la population belgerdquo BullEtinde La Societe Royale Belge DrsquoAnthropologie et de Prehistoire vol75 pp 67ndash86 1965

[22] PW Brand R B Beach and D EThompson ldquoRelative tensionand potential excursion of muscles in the forearm and handrdquoJournal of Hand Surgery vol 6 no 3 pp 210ndash219 1981

[23] F Ankel-Simons Primate Anatomy An Introduction AcademicPress Orlando Fla USA 2000

[24] Y Ichinose H KanehisaM Ito Y Kawakami and T FukunagaldquoRelationship between muscle fiber pennation and force gen-eration capability in Olympic athletesrdquo International Journal ofSports Medicine vol 19 no 8 pp 541ndash546 1998

Submit your manuscripts athttpwwwhindawicom

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Anatomy Research International

PeptidesInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporation httpwwwhindawicom

International Journal of

Volume 2014

Zoology

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Molecular Biology International

GenomicsInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

The Scientific World JournalHindawi Publishing Corporation httpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BioinformaticsAdvances in

Marine BiologyJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Signal TransductionJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BioMed Research International

Evolutionary BiologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Biochemistry Research International

ArchaeaHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Genetics Research International

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Advances in

Virolog y

Hindawi Publishing Corporationhttpwwwhindawicom

Nucleic AcidsJournal of

Volume 2014

Stem CellsInternational

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Enzyme Research

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

International Journal of

Microbiology

Page 6: Research Article Morphometric and Statistical Analysis of ...downloads.hindawi.com/journals/bmri/2014/178906.pdf · frequency or case studies, but comparative anatomy in primates

6 BioMed Research International

followed by PL from primitive to derived primates associatedwith aspects such as the muscular force difference betweenprimates and verification of anatomical aspects in morespecies and consider intra- and interspecific variations

5 Conclusions

Apparently in primates the PL tendon relative size seems toincrease from ancestors genera to more derived ones Thissuggests that this muscle is weaker in terrestrial primateswhen compared to arboreal primates The PL tendon seemsto be more derived in Gorilla sp considering the factorsanalyzed here but the tendon is longer in modern humans

Conflict of Interests

The authors declare that there is no conflict of interestsregarding the publication of this paper

References

[1] S Standring ldquoPelvis girdle and lower limbrdquo in Grayrsquos Anatomythe Anatomical Basis of Clinical Practice Churchill LivingstoneLondon UK 2008

[2] J W M Kigera and S Mukwaya ldquoFrequency of agenesisPalmaris longus through clinical examinationmdashan East Africanstudyrdquo PLoS ONE vol 6 no 12 Article ID e28997 2011

[3] H Gray ldquoMembro superiorrdquo in Gray Anatomia GuanabaraKoogan Rio de Janeiro Brazil 1979

[4] V S Telles and B A L Ernesto ldquoConsideraciones anatomicasde los musculos inconstantesrdquo MedUnab vol 1 no 3 pp 165ndash170 1998

[5] J M Cope E M Looney C A Craig R Gawron R Lamprosand R Mahoney ldquoMedian nerve compression and the reversedpalmaris longusrdquo International Journal od Anatomical Varia-tion vol 2 pp 102ndash104 2009

[6] A B M Machado and L J A Di Dio ldquoFrequency of themusculus palmaris longus studied in vivo in some Amazonindiansrdquo American Journal of Physical Anthropology vol 27 no1 pp 11ndash20 1967

[7] M A Babinski K M A Valente C M S Savedra C H FBurity and A M S Sabra ldquoVariacao anatomica do musculopalmar longo associado a ausencia do arco palmar superficiale compressao do canal de GuyonrdquoActa Scienteae Medica vol 2no 2 pp 54ndash58 2008

[8] C Fowlie C Fuller and M K Pratten ldquoAssessment of thepresenceabsence of the palmaris longus muscle in differentsports and elite andnon-elite sport populationsrdquoPhysiotherapyvol 98 no 2 pp 138ndash142 2012

[9] L Testut and A Latarjet Tratado de Anatomia Humana SalvatEditores Barcelona Espana 1959

[10] S Gibbs Comparative soft tissue morphology of the extantHominoidea including man [PhD thesis] The University ofLiverpool Inglaterra 1999

[11] A O Kayode A A Olamide I O Blessing and O U VictorldquoIncidence of palmaris longus muscle absence in Nigerianpopulationrdquo International Journal of Morphology vol 26 no 2pp 305ndash308 2008

[12] L C Angelini Junior F B Angelini B C Oliveira S ASoares L C Angelini and R H Cabral ldquoUse of the tendon

of the palmaris longus muscle in surgical proedures study oncadaversrdquoActaOrtopedica Brasileira vol 20 no 4 pp 226ndash2292012

[13] T A Aversi-Ferreira L G Vieira R M Pires Z Silvaand N Penha-Silva ldquoEstudo anatomico dos musculos flexoressuperficiais do antebraco no macaco Cebus apellardquo BioscienceJournal vol 22 no 1 pp 139ndash144 2006

[14] T A Aversi-Ferreira ldquoA new statistical method for comparativeanatomyrdquo International Journal of Morphology vol 27 no 4 pp1051ndash1058 2009

[15] T A Aversi-Ferreira R S Maior F O Carneiro-e-Silva et alldquoComparative anatomical analyses of the forearm muscles ofCebus libidinosus (Rylands et al 2000) manipulatory behaviorand tool userdquo PLoS ONE vol 6 no 7 Article ID e22165 8 pages2011

[16] H Preuschoft ldquoMuscles and joints of the anterior extremity ofa gorilla (Gorilla gorilla Savage et Wyman 1847)rdquo Gegenbaursmorphologisches Jahrbuch vol 107 no 2 pp 99ndash183 1965

[17] E E Sarmiento ldquoTerrestrial traits in the hands and feet ofgorillasrdquo American Museum Novitates vol 3091 pp 1ndash56

[18] R Diogo J F Pastor E M Ferrero et al Photographic andDescriptive Musculoskeletal Atlas of Gorilla With Notes on theAttachments Veriations Innervation Synonymy and Weight ofthe Muscles Science Publisher Enfield USA 2010

[19] A Keith ldquoOn the chimpanzees and their relationship to thegorillardquo Proceedings of the Zoological Society of London vol 67no 2 pp 296ndash312 1899

[20] E Loth Anthropologie Des Parties Molles (Muscles IntestinsVaisseaux Nerfs Peripheriques) Mianowski-Masson et CieParis France 1931

[21] G T Meglioli ldquoEtude de Irsquoheredite du muscle petit palmaire(Musculus palmaris longus) dans la population belgerdquo BullEtinde La Societe Royale Belge DrsquoAnthropologie et de Prehistoire vol75 pp 67ndash86 1965

[22] PW Brand R B Beach and D EThompson ldquoRelative tensionand potential excursion of muscles in the forearm and handrdquoJournal of Hand Surgery vol 6 no 3 pp 210ndash219 1981

[23] F Ankel-Simons Primate Anatomy An Introduction AcademicPress Orlando Fla USA 2000

[24] Y Ichinose H KanehisaM Ito Y Kawakami and T FukunagaldquoRelationship between muscle fiber pennation and force gen-eration capability in Olympic athletesrdquo International Journal ofSports Medicine vol 19 no 8 pp 541ndash546 1998

Submit your manuscripts athttpwwwhindawicom

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Anatomy Research International

PeptidesInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporation httpwwwhindawicom

International Journal of

Volume 2014

Zoology

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Molecular Biology International

GenomicsInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

The Scientific World JournalHindawi Publishing Corporation httpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BioinformaticsAdvances in

Marine BiologyJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Signal TransductionJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BioMed Research International

Evolutionary BiologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Biochemistry Research International

ArchaeaHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Genetics Research International

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Advances in

Virolog y

Hindawi Publishing Corporationhttpwwwhindawicom

Nucleic AcidsJournal of

Volume 2014

Stem CellsInternational

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Enzyme Research

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

International Journal of

Microbiology

Page 7: Research Article Morphometric and Statistical Analysis of ...downloads.hindawi.com/journals/bmri/2014/178906.pdf · frequency or case studies, but comparative anatomy in primates

Submit your manuscripts athttpwwwhindawicom

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Anatomy Research International

PeptidesInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporation httpwwwhindawicom

International Journal of

Volume 2014

Zoology

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Molecular Biology International

GenomicsInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

The Scientific World JournalHindawi Publishing Corporation httpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BioinformaticsAdvances in

Marine BiologyJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Signal TransductionJournal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BioMed Research International

Evolutionary BiologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Biochemistry Research International

ArchaeaHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Genetics Research International

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Advances in

Virolog y

Hindawi Publishing Corporationhttpwwwhindawicom

Nucleic AcidsJournal of

Volume 2014

Stem CellsInternational

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Enzyme Research

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

International Journal of

Microbiology