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Accepted manuscript posted online 23 March 2016 , doi: 10.1128/CVI.00721-15

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Controversy exists regarding the optimal use of the 23-valent pneumococcal conjugate vaccine for the protection of high-risk individuals, such as children and adults with immunocompromising conditions and the elderly. The effectiveness and immunogenicity of 23-valent pneumococcal polysaccharide vaccine (PPV23) are limited in such high-risk populations compared to the healthy, with meta-analyses failing to provide robust evidence on vaccine efficacy against invasive pneumococcal disease (IPD) or pneumonia. Moreover, several studies have demonstrated a PPV23-induced state of immune tolerance or hyporesponsiveness to subsequent vaccination, where the response to revaccination does not reach the levels achieved with primary vaccination. The clinical significance of hyporesponsiveness is not yet clarified, but attenuated humoral and cellular response could lead to reduced levels of protection and increased susceptibility to pneumococcal disease. As disease epidemiology among high-risk groups shows that we are still in need of maximum serotype coverage, the optimal use of PPV23 in the context of combined conjugate/polysaccharide vaccine schedules is an important priority. In this minireview, we discuss PPV23-induced hyporesponsiveness and its implications in designing highly effective vaccination schedules for the optimal protection for high-risk individuals.

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S treptococcus pneumoniae (the pneumococcus) is a major cause of life-threatening invasive infections accounting for considerable morbidity and mortality worldwide ( Runner up sneakers Black Camper EcIIH
). Children and adults with certain medical conditions as well as the elderly are at increased risk for invasive pneumococcal disease (IPD) and pneumonia, with disease rates up to 20 times higher than those in the general population ( 2 ). High-risk groups consist of children and adults with chronic diseases and with primary and secondary immune deficiencies and of persons with functional or anatomical asplenia. Persons older than 65 years of age are also at increased risk for pneumococcal infection, due to attenuation of their immune response caused by advancing age, a phenomenon called immunosenescence ( 3 ). The number of patients in need for protection against IPD is continually increasing due to rising numbers of people with chronic disease or HIV infection and an aging population in many high-income countries.

Pneumococcal disease is usually more severe in such high-risk individuals than in immunocompetent subjects ( 4 Womens Granada Closed Toe Heels Jenny 1hIg6
). While antibiotic resistance represents an additional hurdle for the successful treatment of pneumococcal infections ( 7 , 8 ), optimal protection of such “high-risk” groups against S. pneumoniae infection through vaccination continues to be an important priority.

In summary, the continuous large-scale deformation of the wing blade emerges from complex patterns of cell dynamics on small scales. During phase I, cells undergo AP-oriented T1 transitions while elongating in the PD axis. Cell divisions during phase I contribute shear along the PD axis. During phase II, the orientation of T1 transitions shifts to the PD axis and cells relax their shape. Correlation effects contribute AP shear, and peak roughly at the time that T1's shift their orientation.

To ask whether the cellular contributions to tissue shear were independent of each other, we compared the sum of the variances of the final cellular contributions to tissue shear with the variance of final tissue shear itself. The ratio of these values is about 25, indicating that the cellular contributions to tissue shear—like the contributions to area change—are not independent of each other. Thus, the overall tissue shear is more reproducible than would be expected from the variations of the cellular contributions.

What exactly are cells doing that results in correlation effects? We found that shear due to correlation effects was mainly generated by correlations between local elongation and rotations ( Bordaux Velvet Tassel Dress S / RED I Saw It First SyOJEdX
). To investigate this further, we determined the magnitude and orientation of shear and the rotation rate associated with each triangle for each frame of the video. We observed that triangles rotated and sheared in striking spatial patterns that rapidly fluctuate in time ( Figure 6C,F,G ). These patterns correspond to rows of correlated shear and rotation that are distributed throughout the wing blade. To characterize these correlated patterns, we calculated the spatial power spectrum of the local tissue rotation rate ( Womens Messi Espadrilles New Look TERm7
, Mens Slim Jeans Joop GPBkTUQdND
, ‘Power spectrum of local tissue rotation’). This revealed that shear and rotation are correlated in regions corresponding to PD-oriented rows that were about 3–7 triangles long. These rows consist of triangles that all rotate and shear in the same direction. The rows are interspersed with other rows of similar length with mirrored patterns of shear and rotation (note blue and red rows in Womens Simone Dusky Floral Dress Fat Face VzhqS
). Such a pattern of rotation and pure shear is characteristic of neighboring rows of triangles and cells undergoing simple shear in alternating directions ( Womens Mesh Dress Miss Selfridge qZtWoE
). This would occur if PD-oriented rows of cells slide with respect to each other. As discussed above, such rearrangements can indeed contribute to correlation terms (see cartoon in Figure 4H ).

If rows of cells slide past each other, cells typically engage in T1 transitions. Since the peak of AP correlation effects coincided with a shift in the net orientation of T1 transitions (i.e., when the red curve in Figure 6A crosses 0), we wondered whether correlation effects could be associated with T1 transitions at this time (see also Figure 6—figure supplement 1C ). We therefore examined whether correlation effects were associated with a particular type of topological change. Indeed, correlation effects are mostly accounted for by those cells that are about to undergo a T1 transition within the next 9 frames, although they cover only a small fraction of the total area ( Figure 6I , Appendix 1 , ‘Role of T1 transitions in the correlation-induced shear’).

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