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Turkey of the week: Jacob's, Publicis London
James Swift wonders where this Jacob's ad went wrong: "It feels as though there might have been an idea here at some point, but it got diluted along the way. Balls-out weirdness or a funnier execution might have saved the spot but, as it stands, it's neither one thing nor the other."
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Pair of Islanders win opening matches
WACO Texas A&M University-Corpus Christi men's tennis players Mikhail Pavlov and Dmitry Novikov won their first matches at the HEB Baylor Invitational before falling in the second round.
Pavlov defeated Milos Kustudija of Texas Tech, 6-4, 3-6, 6-3 in the opening-round of 64 before falling to No. 1 seed Erling Tveit of Ole Miss, 6-2, 6-1. Novikov, playing in the A draw as well, took down Brad Williams of Texas A&M, 6-4, 5-7, 6-3 before withdrawing from his match against David Simon of Louisville.
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Musculoskeletal System - Limb Development
|Embryology - 28 Nov 2014 Translate
Arabic | Chinese (simplified) | French | German | Hebrew | Hindi | Indonesian | Japanese | Korean | Portuguese | Romanian | Russian | Spanish These external translations are automated and may not be accurate.
- 1 Introduction
- 2 Some Recent Findings
- 3 Textbooks
- 4 Objectives
- 5 Development Overview
- 6 Limb Axis Formation
- 7 Week 5
- 8 Week 6
- 9 Week 7
- 10 Week 8
- 11 Limb Rotation
- 12 Interdigital Apoptosis
- 13 Fetal Growth
- 14 Limb Bone
- 15 Shoulder and Pelvis
- 16 Limb Skeletal Muscle
- 17 Wing as Limb Model
- 18 Mouse Limb Model
- 19 Bat Limb Model
- 20 Molecular
- 21 References
- 22 Additional Images
- 23 External Links
- 24 Glossary Links
The mesoderm forms nearly all the connective tissues of the musculoskeletal system. Each tissue (cartilage, bone, and muscle) goes through many different mechanisms of differentiation.
The musculoskeletal system consists of skeletal muscle, bone, and cartilage and is mainly mesoderm in origin with some neural crest contribution.
The intraembryonic mesoderm can be broken into paraxial, intermediate and lateral mesoderm relative to its midline position. During the 3rd week the paraxial mesoderm forms into "balls" of mesoderm paired either side of the neural groove, called somites.
Somites appear bilaterally as pairs at the same time and form earliest at the cranial (rostral,brain) end of the neural groove and add sequentially at the caudal end. This addition occurs so regularly that embryos are staged according to the number of somites that are present. Different regions of the somite differentiate into dermomyotome (dermal and muscle component) and sclerotome (forms vertebral column). An example of a specialized musculoskeletal structure can be seen in the development of the limbs.
Skeletal muscle forms by fusion of mononucleated myoblasts to form mutinucleated myotubes. Bone is formed through a lengthy process involving ossification of a cartilage formed from mesenchyme. Two main forms of ossification occur in different bones, intramembranous (eg skull) and endochondrial (eg limb long bones) ossification. Ossification continues postnatally, through puberty until mid 20s. Early ossification occurs at the ends of long bones.
Musculoskeletal and limb abnormalities are one of the largest groups of congenital abnormalities.
- Musculoskeletal Links: Introduction | Mesoderm | Somitogenesis | Limb | Cartilage | Bone | Bone Timeline | Axial Skeleton | Skull | Joint | Muscle | Muscle Timeline | Tendon | Diaphragm | Lecture - Musculoskeletal Development | Abnormalities | Limb Abnormalities | Cartilage Histology | Bone Histology | Skeletal Muscle Histology | Category:Musculoskeletal
|Historic Embryology
|1902 Limbs | 1910 Limb Muscles | 1922 Pig Limb Vasculature
- Factor Links: BMP | Sonic hedgehog | Homeobox | FGF | Nanog | Notch | Pax | Retinoic acid | Sox | Tbx | TGF-beta | VEGF | Wnt | hCG
Some Recent Findings
|More recent papers
References listed on the rest of the content page and the associated discussion page (listed under the publication year sub-headings) do include some editorial selection based upon both relevance and availability.
Farnoosh Saraee, Mohsen Sagha, Homa Mohseni Kouchesfehani, Alireza Abdanipour, Masoud Maleki, Mahin Nikougoftar Biological parameters influencing the human umbilical cord-derived mesenchymal stem cells' response to retinoic acid. Biofactors: 2014; PMID: 25408532 Maryna Masyuk, Beate Brand-Saberi Recruitment of Skeletal Muscle Progenitors to Secondary Sites: A Role for CXCR4/SDF-1 Signalling in Skeletal Muscle Development. Results Probl Cell Differ: 2015, 56;1-23 PMID: 25344664 Xin Cheng, Jian-Long Chen, Zheng-Lai Ma, Zhao-Long Zhang, Shun Lv, Dong-Mei Mai, Jia-Jia Liu, Manli Chuai, Kenneth Ka Ho Lee, Chao Wan, Xuesong Yang Biphasic influence of dexamethasone exposure on embryonic vertebrate skeleton development. Toxicol. Appl. Pharmacol.: 2014; PMID: 25286332 Dana Horakova, Petra Cela, Pavel Krejci, Lukas Balek, Simona Moravcova Balkova, Eva Matalova, Marcela Buchtova Effect of FGFR inhibitors on chicken limb development. Dev. Growth Differ.: 2014, 56(8);555-72 PMID: 25280231 G Manivannan, Premal Das, G Karthikeyan, Annamma S John Reconstructive surgery in children to correct ulnar claw hand deformity due to leprosy. Lepr Rev: 2014, 85(2);74-80 PMID: 25255610
Elisa Bettoni, Giorgio Ferriero, Hadeel Bakhsh, Elisabetta Bravini, Giuseppe Massazza, Franco Franchignoni A systematic review of questionnaires to assess patient satisfaction with limb orthoses. Prosthet Orthot Int: 2014; PMID: 25428901 Z Sedaghat, M Kadkhodaee, B Seifi, P Ahghari Hepatoprotective effects of remote perconditioning during renal ischemia. Bratisl Lek Listy: 2014, 115(11);675-679 PMID: 25428534 Zongxi Li, Emmanuel Roussakis, Pieter G L Koolen, Ahmed M S Ibrahim, Kuylhee Kim, Lloyd F Rose, Jesse Wu, Alexander J Nichols, Yunjung Baek, Reginald Birngruber, Gabriela Apiou-Sbirlea, Robina Matyal, Thomas Huang, Rodney Chan, Samuel J Lin, Conor L Evans Non-invasive transdermal two-dimensional mapping of cutaneous oxygenation with a rapid-drying liquid bandage. Biomed Opt Express: 2014, 5(11);3748-3764 PMID: 25426308 J Crawford, K Nygard, B S Gan, D B O'Gorman Periostin induces fibroblast proliferation and myofibroblast persistence in hypertrophic scarring. Exp. Dermatol.: 2014; PMID: 25421393 Agata Paj K, Anna Szczygie, Dorota Paluszy Ska, Barbara Królak-Olejnik Congenital skin aplasia on the lower limb in a premature infant with ELBW ¿ case report. Ital J Pediatr: 2014, 40(1);88 PMID: 25420905
| Keith L. Moore, T.V.N. Persaud, Mark G. Torchia. (2011). The Developing Human: clinically oriented embryology (9th ed.). Philadelphia: Saunders.
| Schoenwolf, G.C., Bleyl, S.B., Brauer, P.R. and Francis-West, P.H. (2009). Larsen's Human Embryology (4th ed.). New York; Edinburgh: Churchill Livingstone Chapter 18 - Development of the Limbs (chapter links only work with a UNSW connection).
- Identify the components of a somite and the adult derivatives of each component.
- Give examples of sites of (a) endochondral and (b) intramembranous ossification and to compare these two processes.
- Identify the general times (a) of formation of primary and (b) of formation of secondary ossification centres, and (c) of fusion of such centres with each other.
- Briefly summarise the development of the limbs.
- Describe the developmental abnormalities responsible for the following malformations: selected growth plate disorders; congenital dislocation of the hip; scoliosis; arthrogryposis; and limb reduction deformities.
Development Overview
Below is a very brief overview using simple figures of 3 aspects of early musculoskeletal development. More detailed overviews are shown on other notes pages Mesoderm and Somite, Vertebral Column, Limb in combination with serial sections and Carnegie images.
Mesoderm Development
Somite Development
|Mesoderm beside the notochord (axial mesoderm, blue) thickens, forming the paraxial mesoderm as a pair of strips along the rostro-caudal axis.
|Paraxial mesoderm towards the rostral end, begins to segment forming the first somite. Somites are then sequentially added caudally. The somitocoel, is a cavity forming in early somites, which is lost as the somite matures.
|Cells in the somite differentiate medially to form the sclerotome (forms vertebral column) and dorsolaterally to form the dermomyotome.
| The dermomyotome then forms the dermotome (forms dermis) and myotome (forms muscle).
Neural crest cells migrate beside and through somite.
| The myotome differentiates to form 2 components dorsally the epimere and ventrally the hypomere, which in turn form epaxial and hypaxial muscles respectively. The bulk of the trunk and limb muscle coming from the Hypaxial mesoderm. Different structures will be contributed depending upon the somite level.
Limb skeletal muscle arises from the hypomere region of the myotomes adjacent to the developing upper (C5-C8) and lower (L3-L5) limb buds.
Limb Axis Formation
Four Concepts - much of the work has been carried out using the chicken and more recently the mouse model of development.
- Limb Initiation
- Proximodistal Axis
- Dorsoventral Axis
- Anteroposterior Axis
Mouse limb Patterning Images
- Mouse Limb Images: Tbx3 and Tbx2 forelimb E10 | Alx3 and Gli3 forelimb E10 | Fgf and Hox forelimb E10.5 | Bmp4 forelimb E11.5 | Bmp4 hindlimb E11.5 | Shh forelimb E11.5 | Fgf8 hindlimb E11.5 | Sox9 forelimb E12.5 | Msx2 forelimb E12.5 | Shh hindlimb E12.5
- Links: Fgf | Hox | Shh | Sox | Limb Development | Mouse Development
Limb Initiation
- Fibroblast growth factor (FGF) coated beads can induce additional limb
- FGF10 , FGF8 (lateral plate intermediate mesoderm) prior to bud formation
- FGF8 (limb ectoderm) FGFR2
- FGF can respecify Hox gene expression (Hox9- limb position)
- Hox could then activate FGF expression
Note that during the embryonic period there is a rostrocaudal (anterior posterior) timing difference between the upper and lower limb development
- this means that developmental changes in the upper limb can precede similar changes in the lower limb (2-5 day difference in timing)
Limb Identity
Forelimb and hindlimb (mouse) identity appears to be regulated by T-box (Tbx) genes, which are a family of transcription factors.
- hindlimb Tbx4 is expressed.
- forelimb Tbx5 is expressed.
- Tbx2 and Tbx3 are expressed in both limbs.
Related Research - PMID: 12490567 | Development 2003 Figures | Scanning electron micrographs of E9 Limb bud wild-type and Tbx5del/del A model for early stages of limb bud growth | PMID: 12736217 | Development 2003 Figures
Tbx3 and Tbx2 expression in E9.75 to 10.5 wild-type mouse embryonic forelimb.[7]
Body Axes
- Anteroposterior - (Rostrocaudal, Craniocaudal, Cephalocaudal) from the head end to opposite end of body or tail.
- Dorsoventral - from the spinal column (back) to belly (front).
- Proximodistal - from the tip of an appendage (distal) to where it joins the body (proximal).
Proximodistal Axis
- Apical Ectodermal Ridge (AER) formed by Wnt7a
- then AER secretes FGF2, 4, 8
- stimulates proliferation and outgrowth
The developing limb can be described along the proximodistal axis as having three main regions:
- Stylopod - the proximal region the limb, the skeletal component of the upper limb (forelimb) is the humerus, and for the lower limb (hindlimb) is the femur.
- Zeugopod - the mid-section of the limb , the skeletal components of the upper limb (forelimb) are the radius and ulna, and for the lower limb (hindlimb) are the tibia and fibula.
- Autopod - the distal region the limb, the musculoskeletal component of the upper limb (forelimb) is the hands, and for the lower limb (hindlimb) is the foot.
Dorsoventral Axis
- Somites - provides dorsal signal to mesenchyme which dorsalizes ectoderm
- Ectoderm - then in turn signals back (Wnt7a) to mesenchyme to pattern limb
- name was derived from 'wingless' and 'int'
- Wnt gene first defined as a protooncogene, int1
- Humans have at least 4 Wnt genes
- Wnt7a gene is at 3p25 encoding a 349aa secreted glycoprotein
- patterning switch with different roles in different tissues
- mechanism of Wnt and receptor distribution still being determined (free diffusion, restricted diffusion and active transport)
One WNT receptor is Frizzled (FZD)
- Frizzled gene family encodes a 7 transmembrane receptor
Fibroblast growth factors (FGF)
- Family of at least 17 secreted proteins
- bind membrane tyrosine kinase receptors
- Patterning switch with many different roles in different tissues
- FGF8 = androgen-induced growth factor, AIGF
FGF receptors
- comprise a family of at least 4 related but individually distinct tyrosine kinase receptors (FGFR1- 4) similar protein structure
- 3 immunoglobulin-like domains in extracellular region
- single membrane spanning segment
- cytoplasmic tyrosine kinase domain
Anteroposterior Axis
- Zone of polarizing activity (ZPA)
- a mesenchymal posterior region of limb
- secretes sonic hedgehog (SHH)
- note digit 1 (thumb/big toe) is the only digit that forms independent of SHH activity.
- apical ectodermal ridge (AER), which has a role in patterning the structures that form within the limb
- majority of cell division (mitosis) occurs just deep to AER in a region known as the progress zone
- A second region at the base of the limbbud beside the body, the zone of polarizing activity (ZPA) has a similar patterning role to the AER, but in determining another axis of the limb
Week 5
|Carnegie stage 13
|Carnegie stage 14
|Carnegie stage 15
- Links: Week 5 | Carnegie stage 13 | Carnegie stage 14 | Carnegie stage 15
Week 6
Digital rays become visible on the upper limb.
- Links: Week 6 | Carnegie stage 16 | Carnegie stage 17
Week 7
|Carnegie stage 18
|Carnegie stage 19
Digital rays become visible on the lower limb.
- Links: Week 7 | Carnegie stage 18 | Carnegie stage 19
Week 8
- Links: Week 8 | Carnegie stage 20 | Carnegie stage 21 | Carnegie stage 22 | Carnegie stage 23
Limb Rotation
Human Embryo (stage 19) showing direction of limb rotation.
Interdigital Apoptosis
Early development of both the hand and foot appear initially as "paddles" at the end of the upper and lower limb respectively. As they continue to grow the digits (fingers and toes) are initially "webbed" together and the cells in the webbing die by programmed cell death to form the separate digits, this process is described as interdigital apoptosis.
Interdigital apoptosis, like general limb growth, occurs first in the upper limb and then later in the lower limb.
- Links: Apoptosis
Fetal Growth
| Embryonic period - the external appearance of both the upper and lower limb has been formed.
Play the associated animation to observe the relative change in limb dimensions.
Limb Bone
|Table Of Ossification Of The Bones Of The Superior Extremity
|(Days and weeks refer to the prenatal, years to the postnatal period.)
|Time of appearance of centre
|Union of primary and secondary centres; remarks.
|6th week
|There are two centres in the shaft, a medial and a lateral. These blend on the 45th day (Mall). Shaft and epiphysis unite between the 20th and 25th years.
|Sternal epiphysis
|18th to 20th year
|Primary centres:
|The chief centre appears near the lateral angle. The subcoracoid centre appears at the base of the coracoid process and also gives rise to a part of the superior margin of the glenoid fossa. The coracoid process joins the body about the age of puberty. The acromial epiphysis centres (two or three in number) fuse with one another soon after their appearance and with the spine between the 22nd and 25th years (Quain); 20th year (Wilms). The subcoracoid and the epiphysis of the coracoid process, the glenoid fossa, the inferior angle, and the vertebral margin join between the 18th and 24th years in the order mentioned (Sappey).
|1. That of the body, the spine, and the base of the glenoid cavity.
|8th week (Mall)2
|2. Goraooid process
|1st year
|3. Subcoracoid
|10th to 12th year
|Acromial epiphyses
|15th to 18th year
|Epiphysis of the inferior angle.
|16 to 18th year
|Epiphyses of the vertebral border.
|18th to 20th year
|Epiphyses of upper surface of coracoid.
|16th to 18th year.
|Epiphysis of surface of glenoid fossa.
|16th to 18th year.
|6th to 7th week (Mall)
|The epiphyses of the head, the tuberculum majus and the tuberculum minus (the last is inconstant) unite with one another in 4th-6th year and with the shaft in 20th-25th year. The epiphyses of the capitulum, lateral epicondyle, and trochlea unite with one another and then in the 16th-17th year join the shaft. The epiphysis of the medial epicondyle joins the shaft in the 18th year.
|1st to 2d year
|Tuberculum majus
|2d to 3d year
|Tuberculum minus
|3d to 5th year
|2d to 3d year
|Epioondylus med
|5th to 8th year
|Lateral margin of trochlea
|11th to 12th year
|Epicondylus lat
|12th to 14th year
|7th week (Mall)
|The superior epiphysis and shaft unite between the 17th and 20th years. The inferior epiphysis and shaft about the 21st year (Pryor); M 21st year, F 21st-25th year (Sappey). Sometimes an epiphysis is found m the tuberosity (R. and K.) and in the styloid process (Sappey).
|Carpal end
|F 8th month - M 15th month (Pryor)
|Humeral end
|6th-7th year
|7th week
|The centre for the shaft of the ulna arises a few days later than that for the radius. The proximal epiphysis is united to the shaft about the 17th year; the inferior epiphysis between the 18th and 20th years; F 20th - 21st years, M 21st - 24th years (Sappey). There is sometimes an epiphysis in the styloid process (Sohwegel) and in the tip of the olecranon process (Sappey).
|Carpal end
|F 6th-7th year - M 7th-8th year (Pryor)
|Humeral end
|10th year
|Os capitatum
|F 3d-6th month M 4th-10th month
|The navicular sometimes has two centres of ossification (Serres. Rambaud and Renault). Serres and Pryor have described two centres of ossification in the lunatum. Debierre has described two centres in the pisiform, one in a girl of eleven, the other in a boy of twelve. The OS hamatum may have a special centre for the hamular process. Pryor has found two centres in the triquetrum. Pryor (1908), describes the centres of ossification of the carpal bones as assuming shapes characteristic of each bone at an early period.
|Os hamatum
|F 5th-10th month M 6th-12th month
|Os triquetrum
|F 2d-3d year M about 3 years
|Os lunatum
|F 3rd-4th year M about 4 years
|Os naviculare
|F at 4 years, or early in 5th year M about 5 years
|Os mult. maj.
|F 4th-5th year M 5th-6th year
|Osmult. min.
|F 4th-5th year M 6th-6th year
|Os pisiforme
|F 9th-10th year M 12th-3th year
|9th week (Mall)
|The centres for the shafts of the second and third metacarpals are the first to appear. There may be a distal epiphysis for the first metacarpal and a proximal epiphysis for the second. Pryor (1906). found the distal epiphysis of the first metacarpal in about 6 per cent, of cases. It is a family characteristic. It arises before the 4th year and unites later. Pryor found the proximal epiphysis of the second metacarpal in six out of two hundred families. It unites with the shaft between the 4th and 6th-7th year; sometimes, however, not until the 14th year. In the seal and some other animals all the metacarpals have proximal and distal epiphyses (Quain). The epiphyses join the shafts between the 15th and 20th years. There may bean independent epiphysis for the styloid process of the 5th metacarpal. The epiphysis of the metacarpal of the index finger appears first. This is followed by those of the 3d, 4th, 5th, and 1st digits.
|Proximal epiphysis of the first metacarpal
|3d year
|Distal epiphyses of the metacarpals
|2d year
|9th week (Mall)
|First row
|Proximal epiphyses
|1st-3rd year (Pryor)
|The shafts of the phalanges of the second and third fingers are the first to show centres of ossification. The phalanges of the little finger are the last, the epiphysis in the middle finger is the first to appear. This is followed by those of the 4th, 2d, 5th, and 1st digits.
|Middle row
|11th-12th week (Mall)
|The centres in the shafts of this row are the last to appear. The epiphysis of the phalanx of the middle finger is the first to appear. This is followed by those of the ring, index, and little finger (Pryor).
|Proximal epiphyses
|2nd-3rd year
|Terminal row
|7th-8th week
|The terminal phalanx of the thumb is the first to show a centre of ossification in the shaft. This is the first centre of ossification in the hand. It is developed in connective tissue while the centres of the other phalanges are developed in cartilage (Mall). The epiphysis of the ungual phalanx of the thumb is followed by those of the middle, ring, index, and little fingers. The fusion of the epiphyses of the phalanges with the diaphyses takes place in the 18th-20th year.
|Proximal epiphyses
|2nd-3rd year
|Sesamoid bones
|Ossification begins generally in the 13th - 14th years, and may not take place until after middle life (Thilenius). For table of relative frequency in the embryo and adult see p. 385.
M = male F = female.
|Table Of Ossification Of The Bones Of The Inferior Extremity
|(Days and weeks refer to the prenatal, years to the postnatal period.)
|Time of appearance of centre
|Time of fusion: general remarks
|Os coxae
|Os ilium
|56th day (Mall)
|The rami of the ischium and the pubis are united by bone in the 7th or 8th year (Quain) ( 12-14 year Sappey). In the acetabulum the three hip bones are separated by a Y-shaped cartilage until after puberty. In this cartilage between the ilium and pubis the "os acetabuli" appears between the ninth and twelfth years. This bone, variable in size, forms a greater or less part of the pubic portion of the articular cavity. Leche (1884). Krause (1885), and many others consider it primarily an independent bone. About puberty between the ilium and ischium and over the acetabular surfaces of these bones small irregular epiphyseal centres appear. The os acetabuli becomes imited to the pubic bone about puberty and soon afterwards the acetabular portions of the ilium and ischium and the ischium and pubis begin to become united by bone. The acetabular portions of the pubis and ilium are unite a little later. Osseous union takes place earlier on the pelvic than on the articular surface of the acetabulum. The union of the several primary centres and the epiphyses is usually completed about the twentieth year.
|Os ischii
|105th day (Mall)
|Os pubis
|4th to 5th fetal month
|Os acetabuli.
|9th to 12th year
| Epiphyses:
Those of the acetabulum
|Soon after puberty
|Crest of ilium
|Soon after puberty
|Fuses with main bone 20th to 25th year
|Tuberosity of ischium
|Soon after puberty
|Fusion begins in the 17th year and is completed between the 20th and 24th years (Sappey)
|Ischial spine
|Soon after puberty
|18th to 20th year (Poirier).
|Ant. inf. spine of ilium
|Soon after puberty
|18th to 20th year (Poirier)
|Symphysis end of os pubis (1 or 2 centres)
|18th to 20th year (Sappey)
|After the 20th year
|43d day (Mall)
| Epiphyses:
Distal end
|Shortly before birth1
|20th to 24th year
|1st year
|18th to 19th year
|Great trochanter
|3d to 4th year (Osseous granules soon after birth, (Poirier)
|18th year
|Small trochanter
| 13th to 14th year
8th year (Sappey)
| 17th year (Quain)
Proximal epiphysis 18th to 22d year (Poirier)
|3d to 5th year
|The osseous patella reaches its definitive form soon before puberty
|44th day (Mall)
| Epiphyses:
Proximal end
|About birth
|19th to 24th year (Sappey)
|Distal end
|2d year
|16th to 19th year
|Tubercle (occas.)
|13th year
|Fuses with epiphysis of the proximal end and then with this to the diaphysis
|55th day (Mall).
| Epiphyses:
Distal end
|2d year
|20th to 22d year
|Proximal end
|3d to 5th year
|22d to 24th year
|Chief centre
|6th fetal month
|The chief nucleus is endochondral. A periosteal nucleus appears frequently in the 4-5 fetal month (Hasselwander)
|Epiphysis (distal end)
| 10th year (Quain)
7th-8th year ( Sappey)
|15th-16th year (Quain)
16th-18th year (Poirier)
M 17-21, average 20 years
F 13-17, average 16 years (Hasselwander)
|6th fetal month (Hasselwander)
|In the 7th-8th year the posterior part of the talus, the os trigonum, is frequently ossified from a special centre (v. Bardeleben). It fuses about the 18th year.
|About birth
|Cuneiform III
|1st year
|Cuneiform I
|2d-3d year
|Cuneiform II
|3d-4th year
|4th-5th year
|8th-10th week
|According to v. Bardeleben a second centre of ossincation appears much later than the primary in the navicular, and finally about the time of puberty a medial epiphyseal centre arises.
|3d-8th year
|The centre for the 2d metatarsal usually appears first, then come the 3rd, 4th, 1st and 5th. The epiphysis of the 1st metatarsal appears at the proximal end of the bone: the other epiphyses arise at the distal ends of the metatarsals. There may be a distal epiphysis in the first metatarsal also.2 In some instances a proximal epiphysis is formed on the tuberosity of the fifth metatarsal (Gruber). The epiphyses unite with the shafts in the 17-21 year in males and in the 14-19 year in females. (Hasselwander).
|Terminal row
|58th day (Mall)
|Epiphyses (distal)
|4th year
| M 13-23, average 16-21 year.
F 13-17, average 14-17 year (Hasselwander).
|Middle row
|4th-10th fetal month
|3d year
| M 15-19 year
F 13-16 year (Hasselwander)
|Proximal row
|3d fetal month
|3d year
| M 15-17 year.
F 14-15 year (Hasselwander)
The centres for the shafts of the phalanges often appear double, one for the dorsal and one for the plantar surface. The centres for the medial phalanges in each row usually appear before the more laterally placed centres. The centre for the 5th terminal phalanx appears much later than the other centres in this row (Mall). According to Rambaud and Renault the epiphyses arise each from two centres which fuse together. In the terminal phalanx of the great toe the ossification centre of the epiphysis often appears as early as the second or even the first year. (Hasselwander)
|Sesamoid bones of the great toe
| M 14th year
F 12th-13th year
|Ossification may begin in the 8th year in females, in the 11th in males (Hasselwander).
M = male F = female.
Bone formation within the limb occurs by endochondral ossification of a pre-existing cartilage template. Ossification then replaces the existing cartilage except in the regions of articulation, where cartilage remains on the surface of the bone within the joint. Therefore bone development in the limb is initially about cartilage development or chondrogenesis.
In addition, there are two quite separate aspects to this development.
- Pattern - where the specific regions will commence to form cartilage, which will be different for each cartilage element.
- Chondrogenesis - the differentiation of mesoderm to form cartilage, which will be essentially the same program for all cartilage templates.
A recent study has identified that the overlying limb surface ectoderm potentially inhibits limb early chondrogenesis through Wnt6 signaling.[12]
Links: Cartilage Development | Bone Development | Bone Development Timeline
Shoulder and Pelvis
The skeletal shoulder consists of: the clavicle (collarbone), the scapula (shoulder blade), and the humerus. Development of his region occurs through both forms of ossification processes.
The skeletal pelvis consists of: the sacrum and coccyx (axial skeleton), and pelvic girdle formed by a pair of hip bones (appendicular skeleton). Before puberty, he pelvic girdle also consists of three unfused bones: the ilium, ischium, and pubis. In chicken, the entire pelvic girdle originates from the somatopleure mesoderm (somite levels 26 to 35) and the ilium, but not of the pubis and ischium, depends on somitic and ectodermal signals.[13]
- Links: Shoulder Development | Pelvis Development
Limb Skeletal Muscle
Embryonic skeletal muscles arise from the somite myotome region giving rise to myogenic progenitor cells.
In the chicken, myotome growth initially occurs by a contribution of myogenic progenitor cells from the medial border of the dermomyotome. Then later by progenitors from all four borders of the dermomyotome, with the medial and the lateral borders of the somite generate the epaxial and hypaxial muscles.[14] Depending on the somite level within the body (trunk), limb or between the limbs, myotome cells will then give rise to limb and trunk muscles respectively.[15]
- limb level somites - cells from the hypaxial (ventrolateral) lips of the dermomyotome delaminate and migrate into the limb bud to form the musculature of the limbs.
- between the limbs somites - cells delaminate from the edges or lips of the epithelial dermomyotome to form the subjacent postmitotic myotome, which gives rise to trunk muscles.
- Early myogenic progenitor cells in the dermomyotome can be initially identified by the transcription factor Pax3.
- Subsequent myogenic program development then depends on the myogenic determination factors (Myf5, MyoD, and MRF4), both Myf5 and MyoD are expressed in the limbs.
- Final differentiation of these cells into post-mitotic muscle fibers in the limb bud is regulated by another myogenic determination factor, Myogenin.
(Some of the above text modified from[15])
- Links: Muscle Development | 1910 Limb Muscles
Wing as Limb Model
- chicken wing easy to manipulate
- removal, addition and rotation of limb regions
- grafting additional AER, ZPA
- implanting growth factor secreting structures
Mouse Limb Model
| Mouse limb skeleton cartoon[16]
Fore-limb and hind-limb buds for stages E9.5 to E13.5. Hindlimbs are morphologically delayed by about half a day.
|Change in cell types and tissue formation as a function of mouse developmental stage.[16]
- Links: Mouse Development
Bat Limb Model
Images of the bat embryo Miniopterus schreibersii fuliginosus at embryonic Stages 13-17.[17]
(aer - apical ectodermal ridge; chp - chiropatagium; eb - elbow; kn - knee)
- Links: Bat Limb Development | Bat Development
Fibroblast Growth Factors
- Fgf8 - morphogen gradient forms by a source-sink mechanism with freely diffusing molecules.[18]
T-box Transcription Factors
The HAND2 gene encodes a basic helix-loop-helix (bHLH).
- Links: Fibroblast Growth Factor | Sonic hedgehog | Wnt | Hand2 | OMIM
- ↑ 1.0 1.1 Anna Kicheva, James Briscoe Limbs made to measure. PLoS Biol.: 2010, 8(7);e1000421 PMID: 20644713 | PMC2903596 | PLoS
- ↑ Alicia Mayeuf-Louchart, Mounia Lagha, Anne Danckaert, Didier Rocancourt, Frederic Relaix, Stéphane D Vincent, Margaret Buckingham Notch regulation of myogenic versus endothelial fates of cells that migrate from the somite to the limb. Proc. Natl. Acad. Sci. U.S.A.: 2014, 111(24);8844-9 PMID: 24927569
- ↑ Elena Kozhemyakina, Andreia Ionescu, Andrew B Lassar GATA6 is a crucial regulator of Shh in the limb bud. PLoS Genet.: 2014, 10(1);e1004072 PMID: 24415953
- ↑ Bea Christen, Alexandre Miguel Cavaco Rodrigues, Monserrat Barragán Monasterio, Carme Fabregat Roig, Juan Carlos Izpisua Belmonte Transient downregulation of Bmp signalling induces extra limbs in vertebrates. Development: 2012, 139(14);2557-65 PMID: 22675213
- ↑ Leila Taher, Nicole M Collette, Deepa Murugesh, Evan Maxwell, Ivan Ovcharenko, Gabriela G Loots Global gene expression analysis of murine limb development. PLoS ONE: 2011, 6(12);e28358 PMID: 22174793
- ↑ Bernd Boehm, Henrik Westerberg, Gaja Lesnicar-Pucko, Sahdia Raja, Michael Rautschka, James Cotterell, Jim Swoger, James Sharpe The role of spatially controlled cell proliferation in limb bud morphogenesis. PLoS Biol.: 2010, 8(7);e1000420 PMID: 20644711 | PMC2903592 | PLoS
- ↑ 7.0 7.1 Antonella Galli, Dimitri Robay, Marco Osterwalder, Xiaozhong Bao, Jean-Denis Bénazet, Muhammad Tariq, Renato Paro, Susan Mackem, Rolf Zeller Distinct roles of Hand2 in initiating polarity and posterior Shh expression during the onset of mouse limb bud development. PLoS Genet.: 2010, 6(4);e1000901 PMID: 20386744
- ↑ Akira Satoh, Aki Makanae, Naoyuki Wada The apical ectodermal ridge (AER) can be re-induced by wounding, wnt-2b, and fgf-10 in the chicken limb bud. Dev. Biol.: 2010, 342(2);157-68 PMID: 20347761
- ↑ Mengyao Dai, Yao Wang, Lu Fang, David M Irwin, Tengteng Zhu, Junpeng Zhang, Shuyi Zhang, Zhe Wang Differential expression of Meis2, Mab21l2 and Tbx3 during limb development associated with diversification of limb morphology in mammals. PLoS ONE: 2014, 9(8);e106100 PMID: 25166052 | PLoS One.
- ↑ Amitabha Bandyopadhyay, Kunikazu Tsuji, Karen Cox, Brian D Harfe, Vicki Rosen, Clifford J Tabin Genetic analysis of the roles of BMP2, BMP4, and BMP7 in limb patterning and skeletogenesis. PLoS Genet.: 2006, 2(12);e216 PMID: 17194222 | PMC1713256 | PLoS Genet.
- ↑ Amitabha Bandyopadhyay, Kunikazu Tsuji, Karen Cox, Brian D Harfe, Vicki Rosen, Clifford J Tabin Genetic analysis of the roles of BMP2, BMP4, and BMP7 in limb patterning and skeletogenesis. PLoS Genet.: 2006, 2(12);e216 PMID: 17194222 | PMC1713256
- ↑ 12.0 12.1 Poongodi Geetha-Loganathan, Suresh Nimmagadda, Bodo Christ, Ruijin Huang, Martin Scaal Ectodermal Wnt6 is an early negative regulator of limb chondrogenesis in the chicken embryo. BMC Dev. Biol.: 2010, 10;32 PMID: 20334703
- ↑ Yegor Malashichev, Bodo Christ, Felicitas Pröls Avian pelvis originates from lateral plate mesoderm and its development requires signals from both ectoderm and paraxial mesoderm. Cell Tissue Res.: 2008, 331(3);595-604 PMID: 18087724
- ↑ Jérôme Gros, Martin Scaal, Christophe Marcelle A two-step mechanism for myotome formation in chick. Dev. Cell: 2004, 6(6);875-82 PMID: 15177035
- ↑ 15.0 15.1 Julien Giordani, Lola Bajard, Josiane Demignon, Philippe Daubas, Margaret Buckingham, Pascal Maire Six proteins regulate the activation of Myf5 expression in embryonic mouse limbs. Proc. Natl. Acad. Sci. U.S.A.: 2007, 104(27);11310-5 PMID: 17592144
- ↑ 16.0 16.1 Leila Taher, Nicole M Collette, Deepa Murugesh, Evan Maxwell, Ivan Ovcharenko, Gabriela G Loots Global gene expression analysis of murine limb development. PLoS ONE: 2011, 6(12);e28358 PMID: 22174793
- ↑ Zhe Wang, Naijian Han, Paul A Racey, Binghua Ru, Guimei He A comparative study of prenatal development in Miniopterus schreibersii fuliginosus, Hipposideros armiger and H. pratti. BMC Dev. Biol.: 2010, 10;10 PMID: 20092640 | PMC: 2824742 | BMC Dev Biol.
- ↑ Shuizi Rachel Yu, Markus Burkhardt, Matthias Nowak, Jonas Ries, Zdenek Petrásek, Steffen Scholpp, Petra Schwille, Michael Brand Fgf8 morphogen gradient forms by a source-sink mechanism with freely diffusing molecules. Nature: 2009, 461(7263);533-6 PMID: 19741606
Justin Cotney, Jing Leng, Jun Yin, Steven K Reilly, Laura E DeMare, Deena Emera, Albert E Ayoub, Pasko Rakic, James P Noonan The evolution of lineage-specific regulatory activities in the human embryonic limb. Cell: 2013, 154(1);185-96 PMID: 23827682
Günter P Wagner, Alexander O Vargas On the nature of thumbs. Genome Biol.: 2008, 9(3);213 PMID: 18341703
Robert E Hill How to make a zone of polarizing activity: insights into limb development via the abnormality preaxial polydactyly. Dev. Growth Differ.: 2007, 49(6);439-48 PMID: 17661738
Akira Satoh, Aki Makanae, Naoyuki Wada The apical ectodermal ridge (AER) can be re-induced by wounding, wnt-2b, and fgf-10 in the chicken limb bud. Dev. Biol.: 2010, 342(2);157-68 PMID: 20347761
Antonella Galli, Dimitri Robay, Marco Osterwalder, Xiaozhong Bao, Jean-Denis Bénazet, Muhammad Tariq, Renato Paro, Susan Mackem, Rolf Zeller Distinct roles of Hand2 in initiating polarity and posterior Shh expression during the onset of mouse limb bud development. PLoS Genet.: 2010, 6(4);e1000901 PMID: 20386744
Emily K Stefanov, Jordan M Ferrage, Nicholas F Parchim, Christine E Lee, Angela D Reginelli, Mara Taché, Rosalie A Anderson Modification of the zone of polarizing activity signal by trypsin. Dev. Growth Differ.: 2009, 51(2);123-33 PMID: 19207183
Search PubMed
Search April 2010
- Limb Development - All (776) Review (108) Free Full Text (196)
- apical ectodermal ridge - All (95) Review (2) Free Full Text (31)
- zone polarizing activity - All (15) Review (2) Free Full Text (7)
Search Pubmed: Limb Development | apical ectodermal ridge | zone polarizing activity | limb bud skeletal muscle
Additional Images
Historic Images
- Limb Images: 274-278 Spinal Column and Lower Limb | 279-284 Lower Limb | 285-288 Knee | 289 Os Coxae | 290 Femur | 291 Tibia | 292 Fibula | 293 Foot | 294 | 295 | 296 | 297 | 298-299 | 300 Forearm and Hand | 301 Upper Limb Joints | 302 Clavicle | Upper Limb Ossification 1 | Upper Limb Ossification 2 | Bone Development Timeline
- Skeleton and Connective Tissues: Connective Tissue Histogenesis | Skeletal Morphogenesis | Chorda Dorsalis | Vertebral Column and Thorax | Limb Skeleton | Skull Hyoid Bone Larynx
Keith, A. (1902) Human Embryology and Morphology. London: Edward Arnold. Chapter 20. The Limbs
External Links
External Links Notice - The dynamic nature of the internet may mean that some of these listed links may no longer function. If the link no longer works search the web with the link text or name.
- Embryo Images - apical ectodermal ridge | AER and vascular channel
- The Jackson Laboratory - Mouse Strains - Limb Patterning Defects
Glossary Links
- A | B | C | D | E | F | G | H | I | J | K | L | M | N | O | P | Q | R | S | T | U | V | W | X | Y | Z | Numbers | Symbols
Cite this page: Hill, M.A. (2014) Embryology Musculoskeletal System - Limb Development. Retrieved November 28, 2014, from https://php.med.unsw.edu.au/embryology/index.php?title=Musculoskeletal_System_-_Limb_Development
- © Dr Mark Hill 2014, UNSW Embryology ISBN: 978 0 7334 2609 4 - UNSW CRICOS Provider Code No. 00098G


text:
1988 Lincoln Town Car1988 Lincoln Town Car Signature Series Sedan model in mint conditionWhite exterior plus a Burgundy-Maroon velour interiorEquipped with a 5.0 Liter... More
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It's not that Fort Lauderdale resident Anni Leoa can't find papayas at her local Publix store. But she thinks they're fresher at Sedano's.
"There is more stuff I like here," said Leoa, who likes to cook recipes from her native Peru. She was among more than 100 shoppers waiting for Broward County's first Sedano's store to open at 8 a.m. Wednesday.


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Edgar Cayce on the Power of Color, Stones, and Crystals
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SF Tidbits for 1/12/07
By John DeNardo | Friday, January 12th, 2007 at 12:23 am
- SCI FI Wire profiles Mark Budz, author of the Philip K. Dick Award finalist Idolon.
- The Sci Phi Show podcast-interviews Mike Resnick.
- Cory Doctorow has started podcasting his novel Eastern Standard Tribe. Be sure to sign up to the RSS feed. [via SFF Audio]
- Eternal Golden Braid ponders the proper reading order for Terry Pratchett's Discworld.
- Friday YouTube: The Ewok Song…as sung by a barbershop quartet. [via The Force.net]
Filed under: Tidbits
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Archives for: June 2012, 23
Turkey threatens action after Syria downs jet
By News on Jun 23, 2012
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text:
But the exhibition group predicts a rebound as the temperatures rise
Average ticket prices topped $8 for the final quarter of the year
The National Association of Theater Owners says movie ticket prices fell four percent over the third quarter
For the first half of the year, the average ticket set back moviegoers $8.02
Convention attendees will get a sneak peak at "The Avengers," "The Dark Knight Rises," and "Men in Black 3" as studios let theater owners know...
Fewer 3D titles meant that ticket prices only jumped four cents in 2011 -- and at the end of the year, they actually got cheaper
Sexually-explicit film could help revive the shunned rating, John Fithian says
It's getting a tiny bit cheaper to see a film, according to the National Association of Theater Owners
Carmike Cinemas CEO S. David Passman, III named chairman of the trade group
The company has now installed almost 8,000 digital screens
Sony said the trade organization is wrong, while shares of RealD plummet 15 percent
The May-through-Labor Day period saw a record $4.4B in ticket sales in the U.S. and Canada, NATO is reporting
John Fithian says studio's early release plan respects windows; Jon Favreau and Quentin Tarantino also oppose move
Hollywood hails legislation that would prevent sites from using U.S. internet service providers, payment processes and search engines
After Jeff Bewkes and Les Moonves say the threat of shorter windows has been overstated, John Fithian begs to differ
"What's an experiment without data?" asks NATO president
NATO data also shows average admissions price declined about 1 percent from the first quarter of 2010
As the controversial premium VOD plan kicks off Thursday, studios are hoping for a longterm payoff. But in the short term, they have nothing but...
Rental platform to launch Thursday with "Just Go With It"
Exhibitors may pull trailers or even films by studios that shorten windows, TheWrap learns
Responding to report that studios are rolling out the new service on DirecTV next month, National Association of Theater Owners puts out an angry statement...
Tension at the annual Las Vegas lovefest exploded on Thursday over a new studio premium VOD plan; exhibitors stressed over drop in box office revenue...
NATO chief John Fithian uses his part of Tuesday's keynote to warn exhibitors: "convert to digital or die"
New MPAA chief will join NATO president and CEO John Fithian to deliver "state of the industry" address on March 29
All those expensive 3D movies at the box office last year yielded one of the biggest year-to-year admissions spikes in history
With studios looking to debut first-run movies in homes days after their theatrical release, exhibitors mobilize in Hollywood and on Wall Street
Price has tumbled 24 cents since peaking at $7.95 in the first quarter
Reflects the fact that more 3D tickets were sold in the first quarter
Average price spiked 6 percent in the first quarter to $7.95, org reports
DGA and NATO join MPAA in asking government regulators to slow down before approving new exchanges


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May 23, 2012 7:46 AM by AP
BATON ROUGE, La. (AP) - A $50 billion, 50-year proposal aimed at stopping coastal land loss in Louisiana has passed the state Legislature.
The Senate voted 94-0 Tuesday in favor of the 2012 coastal master plan. By law, the state has to update it every five years.
Funding for the plan remains uncertain, but the plan envisions getting $50 billion from a variety of sources, including offshore oil and natural gas drilling royalties, federal environmental programs and from what BP PLC may be forced to pay in fines and ecosystem restoration for the 2010 oil spill.
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As I contemplated what to write about in my 50th article for POB, I went back and reviewed my previous articles to see what topics I had covered. I also wondered if there is still a need for a column titled "Back to Basics" considering the changes in technology that have occurred in the 13 years that I have been writing.
The articles I have written over the years have covered a diverse range of topics related to surveying and layout and also some topics that were somewhat philosophical. I have consciously made an effort with every article to select topics that I thought would be of interest to all readers but especially individuals who were just entering the field of surveying. I felt that those who were new to the profession may not be aware of some of the basics of surveying measurement, the fundamentals of instrument use or the standard layout techniques. Fortunately for me, there are always individuals in our profession who can benefit from a review of the basics.
My articles from the past can be separated into a number of distinct categories--including planning, communications, instrument setup and use, calibration, equipment care, mistakes and errors, layout techniques, quick checks, rules of thumb, standard practices--as well as some miscellaneous topics.
One of my favorites was titled "Mistakes and errors from the field" (November 2004) and was based on a questionnaire I sent out to construction layout surveyors as I was working on the 3rd edition of my book, "Construction Surveying and Layout." Just a few of the common mistakes respondents had listed included writing the wrong elevation on a stake and using that elevation to locate a footing; laying out the mirror image of the structure because of lack of understanding of what the plans are communicating; transposing numbers when measuring distance by chaining; using a foot chain and converting to a metric measurement; not holding the prism rod plumb causing the distance to be wrong; not setting up an electronic instrument exactly over the point and not checking to see if the instrument was still level after using it for an extended period of time; not understanding the potential sources of error in total station measurement; failing to turn equal numbers of direct and reverse angles when turning angles; not leveling the instrument when the vertical angle was measured; not "closing the loop when leveling"; not calibrating the instrument before leveling; and being so concerned about the decimal part of the reading when leveling that we read the wrong foot mark.
In "Long backsight and short foresights" (November 2003), I outlined how having a long backsight and a short foresight is a fundamental principle that should always be followed. It is a principle that affects the size of errors in the sighting and angle measurement process of layout. By having a long backsight and a short foresight, the size of sighting errors can be reduced--which is a goal of any good construction layout surveyor. It sometimes isn't convenient and may take more effort to provide a long backsight by requiring someone to walk long distances, but the rewards are great by ensuring that errors will be reduced.
Another favorite of mine was my March 2008 article on "The one-minute, one-person peg test." In this piece, I explained how a peg-test area could be set up using a yardstick, a piece of level rod or a section of carpenter's tape positioned at exactly the same elevation on two stationary objects 200 to 300 feet apart with a clear line of sight between them. Using this peg-test area, one person can perform a calibration test in less than one minute.
Other topics that I have written about over the years include pacing, rules of thumb, quick setup with an optical plummet, leveling an instrument, instrument use, chaining, total stations, lasers in construction, planning, preparation, seminars and education, the role of field engineers, dressing for safety, tolerances, and calculating directions with azimuths.
Oldies but Goodies
Whew, I'm glad I did not have to write all of these articles at one time. It makes me tired to think about all of the effort involved.
I realize some procedures that I have written about are no longer used on a daily basis by many surveyors. I even had someone once say to me, "You're the guy who writes about the old methods." Yep, that's me. Regardless, I do strongly believe that some surveyors and surveyors-in-training still need to hear about some of these old methods of layout and measurement because many of them are still relevant. When the power goes out on the total station or GPS system, knowing some of the "old techniques" may save the day.
I think I can conclude that "Back to Basics" will be applicable for a few more years as long as layout points are put in by people.


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Justin Bieber on SNL: Cute Cameo Alert!by Hilton Hater at . Comments
Justin Bieber completed one of the more packed media blitzes in recent memory this week by appearing on last night's edition of Saturday Night Live.
The 16-year old sensation, teetering dangerously close to overexposure as he promotes the February 11 release of "Never Say Never," stopped by for two segments:
- A skit with host Dana Carvey, in which the comedian reprised his role of "Church Lady" and flirted with the Biebs.
- A digital short with Andy Samberg, in which the pair mocked The Roommate.
Check each out below.
Hey,hey,hey pls the hate of justin bieber dont wst your time writing the commantaire so live him alone i love you justin bieber
USMC just shut the Fuck up no cares if you like him or not MOTHER FUCKING BITCH you can just kill yourself hater
USMC... you just proved Ashley's point. congrats.
Shut the fuck up Ashley, you little bitch!
I don't understand this hatred towards Justin. I think people who spends time hating on him should just a life. The haters are just envious they aren't fortunate as Justin.
Justin you are such a little fucking pussy. If you ever come to my city I'm going to beat the fuck out of you. You are such a crying little fag. I would almost guarantee that you suck cock. You should kill yourself and save the pain that you will have in six months when you're a has been. Fuck you and fuck your mother.
I have always loved snl untill they started having people like Justin beiner on it.... Snl is soposed to be a classic... Justin beiner isn't.
SNL is always hilarious....but i pretty much always have to miss it....either im at a party or sleeping....i would of loved to see yesturdays.....not bcz Justin Wiener but bcz of all of the sketches(or whatever they're called???)


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By Alan Scherstuhl
The Blue Room (La Chambre bleue)
By Alan Scherstuhl
By Stephanie Zacharek
Gone Girl Is Smartly Crafted, Well-Acted, and a Bit Too...
By Stephanie Zacharek
By Aaron Hills
The Subway's the Star in BAM's Transporting Retro
By Melissa Anderson
By Alan Scherstuhl
Terry Gilliam's The Zero Theorem
By Stephanie Zacharek
Even if he's never actually made a film you can stand, you cannot ignore Takashi Miikeno one working today is as preposterously fecund, has visited as many genres (gangster, musical, horror, psychodrama, fantasyoy), or has made each as perversely his own. With this ludicrous fantasy, add children's film to the CVeven if, as the colorful hyperbolists at the New York Asian Film Festival put it, it's a miracle Japanese kids can sleep if this is the kind of film they get. It's really not a shock if you contemplate the state of popness in that manga-anime-deranged culture, but still, it's Miike, chronicling, with as big a budget as he's ever had, the Neverending Storyish saga of a young boy lured into helping out the spirit world against the encroaching forces of robotic technology. The film makes no more or less sense than Ridley Scott's Legend or Jim Henson's Labyrinth, but for many, keeping up with Miike's cranked movie factory output is an end in itself. The DVD includes six making-of featurettes, two freestanding shorts, monster-character profiles, and trailers.
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Vivienne Westwood Small Frilly Snake Crossbody Bag
£195.00 Product Code 3829379
- Overview
A charming Vivienne Westwood classic, this statement Small Frilly Snake Crossbody Bag will take you from the day to evening with ease. Crafted with snake textured faux leather, it features a high-shine finish, flap front and glamorous gold-tone hardware. Complemented with a spacious central compartment and pocket for keeping essentials safe when on the go.
- Details
- Adjustable strap
- Flap front with Vivienne Westwood orb logo
- Popper fastened central compartment with 1 interior zipped pocket
- H11.5cm x W19cm x D4.5cm approx.
- Faux leather
- Includes Vivienne Westwood dust bag
- Delivery & Returns
Delivery Option Duration Price Standard Delivery 3-5 working days, M-F only. £5.95 Next Day Delivery Next working day on all orders placed before 1pm, M-F only. £12 Saturday Delivery Saturday only on orders placed before Friday 1pm £15.95 Delivery to Europe £20 Outside of Europe Click here for International Delivery Terms for overseas shipping. £25
If it's not right, return shipping is free. Find out more.


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WebInsure State
Enabling State-Based Exchanges to create user-friendly, compliant and state-of-the-art online marketplaces
Implementing a state exchange is a daunting task and you will want to work with a provider that has successfully implemented the product before. That company is hCentive. WebInsure State is an ACA-compliant proven exchange that is operational in multiple states.
WebInsure State strictly adheres to CMS guidelines and arrives with standardized, out of the box integration capabilities with state and federal systems. The highly configurable product has UX 2014 compliant interfaces and offers a rich, streamlined user experience. \We are committed to continue to enhance and enrich WebInsure State to ensure compliance, lower cost of ownership, cutting-edge functionality and an intuitive user experience.
WebInsure State is comprised of several portals that are listed below.
Individual Portal: Provides an end-to-end solution for consumers to manage information, eligibility, benefits and enrollments for themselves and their family. In addition, it provides seamless backend integration capabilities for eligibility determination, enrollment and payment transactions.
- Supports intelligent decision-making tools and guided plan shopping for individuals and families allowing the consumer to easily select plans which best meet their health coverage requirements. Our product supports easy eligibility and enrollment renewals thus easing out the burden of.
- Fully compliant to UX 2014 design guidelines, highly configurable and customizable
- Enables consumers with disabilities to perform shopping activities online through an innovative user interface.
SHOP Portal: Designed to assist qualified employers in the enrollment of qualified employees into QHP, including a user-friendly process that makes it easier for small businesses to manage their employee information. The product is a one-stop shop for the employees to search, compare and enroll themselves and their family members in various plan options.
Key Capabilities:
- Provides a friendly interface to submit application for eligibility determination to participate in the State Exchange. The product uses secure and accurate data exchange methods with the external systems for verification.
- Strict adherence to CMS guidelines while enabling high degree of configurability
- Supports multiple contribution methods
- Support management of qualifying life change events for the employees; employees can take appropriate actions to update the enrollment application.
- Multi-employer support
Agent Portal: Allows authorized Agents to assist prospects and customers. The Portal acts as a complete prospect/client management tool and provides agents with all the tools to track and manage their customers and sales pipeline.
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- Enables Brokers to manage sales through seamless automation of clients and proposals management functionalities
- Out-of-the-box capabilities to manage client assignments and sales pipeline including status tracking of sub-agents with pending or completed list of trainings.
- Supports agent affiliation to multi-level agent organization hierarchy and respective certification requirements
- Allows agents actively assist their customers in the process of eligibility and enrollment
Plan Management: SERFF-compliant and enables the State Exchange to perform activities like certifying qualified health plans, collecting and reviewing rate and benefit information, managing plan data before publishing on Exchange, monitoring ongoing compliance issues.
Key Capabilities:
- Allows electronic loading of health insurance plans certified by Department of Insurance (DOI)
- Automates major operational processes such as certification, de-certification of plans, thus allowing exchanges run their operations in an efficient manner
- Java/J2EE & AJAX based application
- Enterprise Service Bus (ESB)/Service Oriented Architecture
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implementation & customization
- Change the look and feel of the application based on your needs
- Integrate with multiple 3rd party systems for analytics/reports/billing needs
- Past successes
- Several U.S. States have selected our product to power their Health Insurance Exchanges.
Frequently Asked Questions
- How does your system conform to the changing federal rules and remains compliant with the PPACA?
- Do you utilize cloud-based environments for your solutions?
Still have questions? Ask our experts
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Systems Administrator1
- Company: Transmara Sugar Company Ltd
- Deadline: Monday 10th October 2011
- Location: Kilgoris
Job Description
Transmara Sugar Company is an upcoming sugar factory and is in the final phase of completion. The factory is situated at Enoosean within Transmara District. The factory is one of its kind, with "state of the art" technology.
- Degree/Diploma in IT
- Should have minimum 3 years experience in a similar position in a busy computing environment.
- Should be conversant with ERP solutions, Networking, Hardware , Software, WiFi, troubleshooting and maintenance
- Should be ready to relocate and work in a rural set up.
How to apply
Interested Candidates who meet the above requirements can send their applications either by Email or Post;
Applications via email should be sent as mentioned;
- CV in MS Word format.
- Application letter with details of current and expected salary.
- All Applicants SHOULD mention the position applied in the Subject of the mail. THIS IS MANDATORY.
- Any email exceeding 1MB will not be considered.
Applications via Post should be sent as mentioned;
- Detailed CV
- Copies of certificates
- ID Copy
- Passport size photos – 2no's
- Any other relevant document.
The applications should be sent to the below address on or before (One week from this advert)
Email: firstname.lastname@example.com
The Human Resource Manager,
Transmara Sugar Company Ltd
P.O.Box- 113, Kilgoris.
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Here's a taste of what TheBody.com has to offer on this topic:
Ask the Experts
Dr. Bob, very worried. Saliva came in touch with open wound,Dr. Bob, An escort was licking my face, later i realised that i had a cut on my face. She could be HIV positive, if she is HIV positive and since her saliva comes into contact with my blood, am i at risk of getting HIV?Let me know. Thanks. I...
Response from Robert J. Frascino, M.D.
The Robert James Frascino AIDS FoundationHi,Nope. Saliva does not transmit HIV. (See below. I'll reprint some basic information about HIV transmission.)Thanks for your tax-deductible donation to The Robert James Frascino AIDS Foundation (www.concertedeffort.org).Stay well.Dr. BobDr.... Read more »
Could I get aids through sweat touching an open woundSo after a game one of my friends drops his cup (for sports) next to me and pick it up quickly... Read more »
"Risk from getting open wound and fingerprick touched"Hi there. I just tested negative ytd 3 months post an unprotected sexual exposure. What I did was... Read more »
San Francisco, Calif., Mayor Lee Celebrates Opening of City's New Bridge HIV Research FacilitySan Francisco Mayor Edwin M. Lee officially cut the ribbon to open the San Francisco Department of Public Health's (DPH) new Bridge HIV research facility, which is part of San Francisco's new $9.5 million HIV/AIDS Research Center. "The research that... Read more »
This Positive Life: Marco Benjamin Lives Openly -- And On the Open RoadIf you have seen an 18-wheel truck with a huge condom on the side driving through your town, then you may have seen Marco Benjamin. Diagnosed with HIV at 27 while working at an architectural firm, Marco left that job within a year to pursue a life... Read more »
Circumcision Alters Penis Microbiome, Could Explain HIV ProtectionCircumcision drastically alters the microbiome of the penis, changes that could explain why circumcision offers protection against HIV and other viral infections. In a study to be published on April 16 in mBio ®, the online open-access journal of... Read more »
Thank You So Much for Thinking About My Penis and Anus -- Online HIV/AIDS Prevention That Focuses on Something That MattersJason Chiu, M.P.H. As a 26-year-old former Grindr-user and a single-and-ready-to-mingle gay man, I am a frequent target of Internet HIV prevention messages/advertisements. Every once in awhile, I do feel obliged to click on the ads because of... Read more »
Connect With Others
if tip of penis just touches opening of penis???
Posted by Anonymous, 1 ReplyIf the tip of my penis went between someones crack and touched the opening of the anal wall without a condom, is it possible to get hiv from this, and if so, what are the odds of it happening. I am circumsized and live in oklahoma, if that has... Read more »
Risk of exposure from open wound and saliva
Posted by mantle198, 2 RepliesHi I'm posting to get a better understanding of my risk exposure and gain some peace of mind, until I wait the 3 month period before I take the HIV test. Recently I had a one night stand with a sex worker, she gave me unprotected oral sex and... Read more »
What's the definition of open wound for HIV
Posted by sweetangel, 1 ReplyWhat's the definition of open wound when it comes to HIV transmission. Does it have to be bleeding? Or how recent to be considered open?
I had sex with my HIV + partner yesterday, He came on my chest but little did I realize that that previous night... Read more »
Hiv transmission through contact of vaginal fluid with an open wound?
Posted by neibet, 1 ReplyWhat is the risk of HIV transmission through the exposure of an open wound to vaginal secretions.
Assuming i am having a back massage and i have an open injury on my back and the masseuse was sitting naked in my back with her vagina in direct... Read more »
Guides for People With HIV


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Here's the scenario....
User logs onto an XP PC, (W2K Server, roaming profiles, re-directed "My Documents") and accesses files in My Documents. The user then deletes a file. The file has gone when you look at the user's folder on the server however, when they log off, the file magically re-appears. I know this is great for stopping some one accidentally deleting files but there are times when a file just needs to be deleted!!!.
All the permissions look OK, user has full control of their home drive down the tree etc.
Any Ideas???
Alan :?


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Code Red Safety Jobs in Indiana
1 Code Red Safety jobs in Indiana found on Monster.Jobs 1 to 1 of 1
Customer Service Manager Code Red Safety is looking for a Customer Service Manager to work at our Corporate Office in Hammond, IN. - (just 34 mins from downtown Chicago) JOB SUMMARY Responsible for overseeing the customer service department and ensuring the company or corporation delivers the highest level of customer service possible. Will supervise, train, coach, and mentor employees. PRIMAR...
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333 Natural Eau de Parfum
Three types of chamomile, three types of lavender, three types of citrus: 333 is a stunning pastiche of some of our favorite botanicals.
Size: 30mL/1fl.oz
Country: United States
Three types of chamomile, three types of lavender, three types of citrus: 333 is a stunning pastiche of some of our favorite botanicals.
More about this product
Daring in its intent, exquisite in its execution, it's an elegantly unique fragrance that stands alone in its aspirations.
Average rating: 4.5 out of 5 hearts
By Bronte on April 27, 2012
Perfect for people with perfume sensitivities
So fresh! I use this daily. Great combination of botanical essences - and no harsh chemicals to trigger my usual perfume sensitivities.
By Naomi on February 15, 2012
Beautiful Fresh Fragrance!
This is a gorgeous and fresh fragrance! The notes of lavender are subtle, and the chamomile/citrus forms the base of the fragrance. This perfume is not floral and is almost unisex...my husband loves it too! I love this perfume for the summer and spring!
Directions for use
Unlike its synthetic cousins, natural perfume doesn't need chemicals to express itself. But it does need a little time to breathe. Give it a minute or two after applying for full effect.
Ethyl Alcohol (Organic Grape Alcohol), Ricinus communis (Organic Castor Oil), Parfum (Fragrance)* *100% Organic, wildcrafted or natural essential oils.


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Dover - CalaisroutesHull - RotterdamroutesHull - ZeebruggeroutesCairnryan - LarneroutesDublin - Liverpoolroutestravel informationroutesTroon - Larneroutes
Dover - CalaisonboardHull - RotterdamonboardHull - ZeebruggeonboardCairnryan - Larne onboardLiverpool DublinonboardIrish Sea improvementsonboardshoppingonboardforeign exchangeonboardtaking your petonboard
short sea offersoffersIrish Sea offersoffersdisneyland offersoffersclub lounge on the Irish Seaoffersseason ticketsoffersFamily Foot passenger Day Tripoffers
2 for 1 minicruises from Hullminicruiseminicruises from Doverminicruisethemed minicruisesminicruiseNew Year's Eve party 2014minicruise
disneyland parisaccommodationsuperbreakaccommodationcottages - France, Ireland, UKaccommodationholiday parksaccommodationluxury mobiles & campingaccommodationskiaccommodationbooking.comaccommodation
BelgiumdestinationsFrancedestinationsHollanddestinationsIrelanddestinationsNorthern IrelanddestinationsScotlanddestinationsSpaindestinationsinspirationdestinations
Dover - Calais
If you are taking your pet(s) on the Dover-Calais route you will need to check-in at least one hour before departure and the animals must remain inside the vehicle at all times during the crossing. Foot passengers cannot travel with pets.
Hull - Rotterdam/Zeebrugge
For the Hull-Rotterdam/Zeebrugge routes you will need to check-in at least ninety minutes before departure and the animals remain inside air-conditioned kennels on the vehicle decks during the duration of the crossing. Coach passengers are unable to carry pets, with the exception of registered guide and hearing dogs.
Liverpool - Dublin, Cairnryan/Troon - Larne
If you are travelling with a pet this must be declared at the time of booking and in advance of travel. Arrangements for pets on board and check-in times vary by route and will be advised at the time of booking. Pets (excluding guide dogs) will not be allowed in the passenger areas of the ship. It is your responsibility to provide all necessary documents and make all arrangements, including where necessary quarantine, to satisfy all legal requirements. Failure to do so will prevent your pet from being permitted to travel. Passengers travelling by coach on any of the P&O Ferries routes and foot passengers using our Irish Sea services will not be permitted to travel with pets. Unaccompanied pets will not be accepted on any of our services. In some cases the carriage of animals will require the payment of freight tariffs. Access to your pet during the crossing is at the discretion of the ships' officers.
We make a charge of £15 each way for each pet travelling onboard Dover to Calais, and £17 for each pet travelling onboard our Hull - Rotterdam/Zeebrugge route, with the exception of registered guide and hearing dogs, which are carried free of charge. For customers travelling on our Irish routes there is no charge for pets.
Please note that animals cannot travel unaccompanied.
PETS passport scheme
For those wishing their dog, cat or ferret to travel under the PETS passport scheme, the most up to date information can be obtained from the Defra website, as well as quarantine requirements for pets that do not qualify under the scheme. Please note, cats and dogs must be vaccinated against rabies. The length of the waiting period before entry to the UK is 21 days after the first vaccination date.
EU legislation now allows rabbits and rodents such as guinea pigs and mice to travel too – and you won't need a pet passport to transport them!
On the Hull-Rotterdam/Zeebrugge route (both foot passenger and motorist bookings) foot passenger bookings cannot be made online so please call us to make your booking.
Always check the Defra website for the most up to date information as P&O Ferries cannot be held responsible for incorrect information on this page.


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En Español
What nipple problems may I have while I am breastfeeding? You may have sore nipples during the first 1 to 2 weeks of breastfeeding. Your nipple and areola (the dark area around your nipple) may look red and feel sore. After the first week of breastfeeding, this discomfort should decrease. It should not hurt to breastfeed. It is never normal for your nipples to bruise or get blisters. Your nipples should not crack, scab, or bleed.
What causes nipple soreness?
What conditions can lead to nipple soreness?
How can nipple soreness affect breastfeeding? You may feel tense if it hurts to breastfeed. This tense feeling may prevent milk from traveling to your nipple (milk let-down). If your baby needs to suck harder to get milk, this can make your nipple pain worse. Do not wait to get help if your nipples are cracked, sore, or painful. You may want to stop breastfeeding when you have sore nipples. Caregivers recommend breastfeeding for at least 6 months.
How can I manage nipple soreness?
How can I prevent nipple soreness?
What should I know about having a pierced nipple and breastfeeding? Remove your jewelry before breastfeeding. Nipple jewelry is dangerous for a breastfeeding baby. Your baby may swallow your jewelry or hurt his mouth or lips on it. Tell your caregiver if your pierced nipple hurts while you breastfeed.
When should I schedule a follow-up visit with my caregiver? Follow up with your caregiver as directed. Write down your questions so you remember to ask them during your visits. Your caregiver may suggest that you see a lactation consultant. This is a caregiver who can help you with breastfeeding.
Where can I get more information?
When should I contact my caregiver? Contact your caregiver if:
© 2014 Truven Health Analytics Inc. Information is for End User's use only and may not be sold, redistributed or otherwise used for commercial purposes. All illustrations and images included in CareNotes® are the copyrighted property of A.D.A.M., Inc. or Truven Health Analytics.
References and sources


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The Last High Ground
By Robin A. White
I know why our aircraft are going down reads the cryptic note left by a disgraced employee of America's leading aircraft manufacturer. But for the passengers and crew of Flight 262, a brand-new aircraft, it comes too late. The worst aviation disaster in U.S. history is about to take place.


text:
I usually get asked about my home-offices. Apparently, people seem to like them and ask for tips on how to create their own. In order to address this, I wrote a huge article about office design back in October 2011 using the story to build my workplace when I was working from home at IBM.
Since then, a lot has changed. Technology evolved a lot in the past 3 years and today my home-office is completely different. So, I thought that an update for that article is adequate. What I am going to show here is valid not only for home-offices, but for traditional offices as well.


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- Frances Holmes on Can Pat Panetta dance? Amanda may hate the hands, but unfortunately Pat that's all you've got moving! more
- Pedrao on Can Pat Panetta dance? You dance like a typical white boy. As a youth, you'de have no chance on scoring. As an old man; who cares. more
- Brad on Can Pat Panetta dance? He looks to enjoying himself and looks natural doing it, so who cares if its good or bad dancing. hahahaCheers Brad ... more
- anna on Can Pat Panetta dance? Pat, no you can't dance. We would like to here the music tho so we can see if your in beat more
- Jude on Can Pat Panetta dance? NO Pat - you cant dance - you dance like my husband!!!!:( :( :( more
- H on Dami Im joins Denis Walter Thanks for having Dami on your show. That was a very good interview, Good questions, Good answers. 3AW, please play her ... more
- kerrie on Denis Walter live from New York Hi Dennis went too new york may 2013 with husband had a great time there check out ellens star dust diner and visit the soup ... more
- Tim on Denis Walter live from New York No wonder that lady named Judy didn't recognise the song you sang to her! She may have if you used the correct words Denis! ... more
- Noel on Denis Walter live from New York Great listening to you in New York.I am reliving my visit and taking note of the sights you are seeing and I hope to see, ... more
- Roger on Denis Walter live from New York Dennis I rang you on live radio about the New York Yacht Club next door to the Sofitel but if you get down to Wall St ... more
- di padgett on Denis Walter live from New York go have a baileys milkshake at Bills Burgers Rockerfellar and do the underground tour of rockerfellar more
Denis Walter's 2012 Carol events
Bringing together families and friends to enjoy an evening of Christmas celebrations!
Here are the details on Denis Walter's two free community concerts.
Saturday December 15 at Hanging Rock
Pre Concert from 7pm with the main concert kicking off at 8pm (Gates open at 5pm)
Guests include international violinist Sally Cooper, rock music identity Wendy Stapleton, Humphrey B. Bear for the kids and, ahead of another overseas tour, The Australian Youth Band. Santa Claus will also be 'Rocking at the Rock'.
Parking will be provided for a gold coin donation, with all funds raised going to the Woodend SES who will be coordinating the car parking at the event.
For further information please email firstname.lastname@example.com
Saturday December 8 at Waterfront Geelong - Eastern Beach Reserve
Concert 7:45pm - 9:30pm
Guests include Julie Walter, Ginger and Tonic, international violinist Sally Cooper, Humphrey B. Bear for the kids and, ahead of another overseas tour, The Australian Youth Band.
Santa Claus will also make a special apperarence!
Denis Walter's Afternoons page
3AW Afternoons provides a feast of entertainment and helpful information between 12 and 4pm every Monday to Friday. Denis Walter, one of 3AW's favourites, hosts the program each day - with highlights regularly uploaded to 3AW.com.au.


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Spam Arrest Loses LawsuitWritten by Sue Walsh on September 27, 2013
Spam filtering service Spam Arrest has lost their lawsuit against a marketing company it claims agreed to their terms of service and then broke them. The company offers a white-listing service as a spam filter. It intercepts unfamiliar emails and demands the sender complete a CAPTCHA if they want their message to be received. The sender is also required to agree to Spam Arrest's terms of service which prohibit unsolicited commercial email (Anyone who has ever sent an email to an Earthlink member is probably familiar with the process). Once the sender complies, their email address is whitelisted and they can email the recipient freely in the future. This type of service is looked on rather dimly by security experts because those who use them seldom bother to whitelist people they correspond with regularly, forcing them to jump through the Spam Arrest hoops, and furthermore when they contact a company's customer service or tech support email, they don't bother to whitelist that email address either. The bottom line is people shouldn't have to jump through hoops to email you.
Getting back to the lawsuit, Spam Arrest sued marketing firm Sentient Jet, saying the company agreed to the terms of service and then proceed to spam their users with over 600 advertising emails. They demanded $2,000 per spam message claiming breach of contract. A U.S. District Court Judge disagreed and threw the suit out, saying Spam Arrest could not prove that Sentient Jet had entered into a legally binding contract. It had no paper trail or signatures.
Neither company has commented on the decision. What do you think? Was Spam Arrest in the right or way out of line?
Related Posts
Shouldn't a court case involve papers and evidence? Did Spam Arrest and their lawyer provide the required evidence? Did they have proof that Sentient Jet really entered into an agreement with them? If they had, then I'd wonder why the District Court Judge threw the case away. On the other hand, a lot of marketing companies use clever strategies and tactics that make their work do not appear like spam. Maybe that's what happened. If this was the case, then only spam experts and insiders will be able to point out the difference between what's legible and what's not. Not all court personnel are familiar with how spammers work. That's just my opinion, though.


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|Home » Crystal Coast » Vacation Rentals » Planning » Rental Gear
|Bogue Banks Beach Gear
Rental Gear
Bluewater offers a variety of beach gear that can be rented for your Crystal Coast vacation! So, forget about loading up the car with all the beach bikes, beach chairs and strapping that kayak to the roof - just rent all your beach essentials through Bluewater. We will make it even easier for you by delivering and picking-up all your rental items for FREE. Weekly rental gear items available for your vacation and prices are listed below. Call (866) 937-0608 and speak with our Vacation Planners today!
Miscellaneous Items:
Forget something? Or didnt have room in the car? Call (866) 937-0608 and speak with our Vacation Planners to schedule delivery. Delivery and Pickup are always FREE!
Remember: Beach gear of any kind cannot be left on the beach overnight. This includes tents, canopies, chairs, nets, umbrellas, etc. Any items left unattended after 7pm will be confiscated by beach patrol. Please be advised that guests are responsible for all confiscated rental gear.
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