Normalized gait analysis parameters are closely related to patient-reported outcome measures after total knee arthroplasty.

Up till now, only a weak connection could be shown between patient-related outcome measures (PROMs) and measurements obtained by gait analysis (e.g. speed, step length, cadence, ground reaction force, joint moments and ranges of motion) after total knee arthroplasty (TKA). This may result from the methodical problem that regression analyses are performed using data that are not normalized against a healthy population. It does appear reasonable to assume that patients presenting a physiological gait pattern are content with their joint. The more the gait parameters differ from a normal gait pattern the worse the clinical outcome measured by PROMs should be expected to be. In this retrospective study, 40 patients were enrolled who had received a gait analysis after TKA, and whose PROMs had been evaluated. A gender- and age-matched control group was formed out of a group of test persons who had already undergone gait analysis. Gait analysis was undertaken using the motion analysis system 3D Vicon with ten infrared cameras and three strength measuring force plates. The physiological gait analysis parameters were deduced from arithmetic mean values taken from all control patients. The deviances of the operated patients' gait analysis parameters from the arithmetic mean values were squared. From these values, the Pearson correlation coefficients for different PROMs were then calculated, and regression analyses were performed to elucidate the correlation between the different PROMs and gait parameters. In the regression analysis, the normalized cadence, relative gait speed of the non-operated side, and range of the relative knee moment of the operated side could be identified as factors which influence the Forgotten Joint Score (FJS-12). The explanation model showed an increase of the FJS-12 with minimisation of these normalized values corresponding to an approximation of the gait pattern seen in the healthy control group. The connection was strong, having a correlation coefficient of 0.708. A physiological gait pattern after TKA results in better PROMs, especially the FJS-12, than a non-physiological gait pattern does.

Effects on proprioception by Kinesio taping of the knee after anterior cruciate ligament rupture.

INTRODUCTION: The use of Kinesio tape (KT) to improve proprioception is a matter of considerable debate. In comparison, the rupture of the anterior cruciate ligament is a sufficiently well-investigated injury with a proven compromise of proprioception. The objective of the present study was to assess a supportive effect on proprioception after KT application, taking the anterior cruciate ligament (ACL) rupture as an example. MATERIALS AND METHODS: Forty-eight patients who had suffered an ACL rupture, confirmed clinically and by magnetic resonance imaging, and who were treated conservatively or were awaiting surgery were included in this study. In all patients, a gait analysis was performed on the affected leg before and after KT application. In addition, the IKDC score, the Lysholm score, stability using the Rolimeter, and the angle reproduction test were determined. RESULTS: Thirty-nine men and nine women who had had an ACL rupture for at least 3 weeks were included in the study. Significant improvements were achieved on the affected knee joint for the gait analysis parameters touchdown and unrolling, cadence, stability and stance phase as well as an extension of the hip joint. The Lysholm score improved from 79.3 to 85.8 (p < 0.001) and the IKDC score from 60.2 to 71.3 points (p < 0.001). Significant improvements were achieved in the Rolimeter and angle reproduction test. CONCLUSIONS: The use of KT has a positive effect on proprioception in patients with an anterior cruciate ligament rupture. Therefore, the application may improve gait pattern as well as the subjective function of the affected knee joint.

Patient-Specific Contact Stress Does Not Predict Polyethylene Wear Rate in a Specific Pressfit Cup.

BACKGROUND: The most common reason for revision total hip arthroplasty remains polyethylene wear. Development dysplasia of the hip and revision situations requires a conscious compromise of implant position. The surgeon should know about the consequence on wear via a possible change in hip contact force. The objective of this study is to investigate whether annual wear is dependent on hip contact force. METHODS: Forty-five inserts (DuraLoc, DePuy) that were explanted in our department were included. Three-dimensional gravimetric determination of the wear was performed by fluid displacement. Then, the hip contact force was determined using radiographs according to the Blumentritt model. RESULTS: No correlation was found between patient-specific factors and the annual wear. The hip contact force estimated by the Blumentritt model also showed no correlation between hip contact force and annual wear. Two single model parameters correlated significantly with wear: VRECAB as a ratio of the lever length of the spinocrural and the pelvitrochanteric muscles and the angle Alpha as a measure of the position of the center of rotation in relation to the greater trochanter. The greater the ratio spinocrural/pelvitrochanteric lever arm (R = 0.408, P = .005) and the greater the Alpha angle (more valgus the femoral neck) (R = 0.377, P = .011) were, the greater was the wear. CONCLUSION: These results lead to the conclusion that neither patient-specific factors nor the estimated hip contact force have a major influence on annual wear in the case of DuraLoc cups. Only a coxa valga and a small femoral offset contribute in a limited amount to an increase in wear.

Forgivingness of an Anteromedially Positioned Small Locked Plate for High Tibial Osteotomy in Case of Overcorrection and Lateral Hinge Fracture.

High tibial osteotomy (HTO) represents a sensible treatment option for patients with moderate unicondylar osteoarthritis of the knee and extraarticular malalignment. The possibility of a continuously variable correction setting and a surgical approach low in complications has meant that the medial opening osteotomy has prevailed over the past decades. The objective of the present study was to determine whether anteromedially positioned small plates are nevertheless forgiving under biomechanically unfavourable conditions (overcorrection and lateral hinge fracture). In this study, a simulated HTO was performed on composite tibiae with a 10-mm wedge and fixed-angle anteromedial osteosynthesis with a small implant. Force was applied axially in a neutral mechanical axis, a slight and a marked overcorrection into valgus, with and without a lateral hinge fracture in each case. At the same time, a physiological gait with a dual-peak force profile and a peak load of 2.4 kN was simulated. Interfragmentary motion and rigidity were determined. The rigidity of the osteosynthesis increased over the cycles investigated. A slight overcorrection into valgus led to the lowest interfragmentary motion, compared with pronounced valgisation and neutral alignment. A lateral hinge fracture led to a significant decrease in rigidity and increase in interfragmentary motion. However, in no case was the limit of 1 mm interfragmentary motion critical for osteotomy healing exceeded. The degree of correction of the leg axis, and the presence of a lateral hinge fracture, have an influence on rigidity and interfragmentary motion. From a mechanically neutral axis ranging up to pronounced overcorrection, the implant investigated offers sufficient stability to allow healing of the osteotomy, even if a lateral hinge fracture is present.

Temporal and spatial relationship between gluteal muscle Surface EMG activity and the vertical component of the ground reaction force during walking.

BACKGROUND: Optimized temporal and spatial activation of the gluteal intermuscular functional unit is essential for steady gait and minimized joint loading. RESEARCH QUESTION: To analyze the temporal relationship between spatially resolved surface EMG (SEMG) of the gluteal region and the corresponding ground reaction force (GRF). METHODS: Healthy adults (29♀; 25♂; age 62.6±7.0 years) walked at their self-selected slow, normal, and fast walking speeds on a 10 m walkway (ten trials/speed). Bilateral paired eight-electrode strips were horizontally aligned at mid-distance of the vertical line between greater trochanter and iliac crest. Concerning the ventral to dorsal direction, the center of each strip was placed on this vertical line. Initially, these signals were monopolarly sampled, but eight vertically oriented bipolar channels covering the whole gluteal region from ventral to dorsal (P1 to P8) were subsequently calculated by subtracting the signals of the corresponding electrodes of each electrode strip for both sides of the body. Three vertical bipolar channels represented the tensor fasciae latae (TFL; P2), gluteus medius (Gmed, SENIAM position; average of P4 and P5), and gluteus maximus muscles (Gmax; P7). To determine the interval between SEMG and corresponding GRF, the time delay (TD) between the respective first amplitude peaks (F1) in SEMG and vertical GRF curves was calculated. RESULTS: Throughout the grand averaged SEMG curves, the absolute amplitudes significantly differed among the three walking speeds at all electrode positions, with the amplitude of the F1 peak significantly increasing with increasing speed. In addition, when normalized to slow, the relative SEMG amplitude differences at the individual electrode positions showed an impressively homogeneous pattern. In both vertical GRF and all electrode SEMGs, the F1 peak occurred significantly earlier with increasing speed. Also, the TD between SEMG and vertical GRF F1 peaks significantly decreased with increasing speed. Concerning spatial activation, the TD between the respective F1 peaks in the SEMG and vertical GRF was significantly shorter for the ventral TFL position than the dorsal Gmed and Gmax positions, showing that the SEMG F1 peak during this initial phase of the gait cycle occurred earlier in the dorsal positions, and thus implying that the occurrence of the SEMG F1 peak proceeded from dorsal to ventral. SIGNIFICANCE: Tightly regulated spatial and temporal activation of the gluteal intermuscular functional unit, which includes both speed- and position-dependent mechanisms, seems to be an essential requirement for a functionally optimized, steady gait.

Mobile Robot-Based Gait Training after Total Hip Arthroplasty (THA) Improves Walking in Biomechanical Gait Analysis.

There are multiple attempts to decrease costs in the healthcare system while maintaining a high treatment quality. Digital therapies receive increasing attention in clinical practice, mainly relating to home-based exercises supported by mobile devices, eventually in combination with wearable sensors. The aim of this study was to determine if patients following total hip arthroplasty (THA) could benefit from gait training on crutches conducted by a mobile robot in a clinical setting. METHOD: This clinical trial was conducted with 30 patients following total hip arthroplasty. Fifteen patients received the conventional physiotherapy program in the clinic (including 5 min of gait training supported by a physiotherapist). The intervention group of 15 patients passed the same standard physiotherapy program, but the 5-min gait training supported by a physiotherapist was replaced by 2 × 5 min of gait training conducted by the robot. Length of stay of the patients was set to five days. Biomechanical gait parameters of the patients were assessed pre-surgery and upon patient discharge. RESULTS: While before surgery no significant difference in gait parameters was existent, patients from the intervention group showed a significant higher absolute walking speed (0.83 vs. 0.65 m/s, p = 0.029), higher relative walking speed (0.2 vs. 0.16 m/s, p = 0.043) or shorter relative cycle time (3.35 vs. 3.68 s, p = 0.041) than the patients from the control group. CONCLUSION: The significant higher walking speed of patients indicates that such robot-based gait training on crutches may shorten length of stay (LOS) in acute clinics. However, the number of patients involved was rather small, thus calling for further studies.

Extremely small genomes in two unrelated dipteran insects with shared early developmental traits.

We discovered extremely small genomes (1C ~100 Mb) in the dipteran insects Coboldia fuscipes (Scatopsidae) and Psychoda cinerea (Psychodidae). The small genomes of these species cannot be explained by a fast developmental rate, which has been shown to correlate with small genome sizes in animals and plants but might accommodate the combined effects of other developmental traits, including small egg size, thin blastoderm layer, and long-germ development.

Systematic differences of gluteal muscle activation during overground and treadmill walking in healthy older adults.

Guteal muscle activation during walkway and treadmill walking was compared by means of Surface EMG (SEMG). Healthy older adults (50-75 years, n = 54; 29 females, 25 males) walked on a walkway (WW) at their self-selected slow, normal, and fast walking speeds and on a treadmill (TM) at 2, 3, 4, 5, and 6 km/h. Subject-individual, best-matched speed pairs were constituted and named SLOW, NORMAL, and FAST. Hip muscle activation was measured on both sides at mid-distance between the greater trochanter and the iliac crest by applying eight equally-spaced bipolar SEMG channels from ventral to dorsal (P1-P8). Grand averaged amplitude curves and mean amplitudes over the complete stride were analyzed to compare WW and TM walking. TM walking evoked significantly elevated mean amplitude levels, particularly at the ventral positions P1 to P4, which were disproportionately increased at SLOW. In grand averaged curves, corresponding significant amplitude differences between WW and TM were observed during load acceptance (SLOW; NORMAL), mid-stance (all speeds), and late swing phase (SLOW), with the number of significant differences decreasing for all electrode positions from SLOW to FAST. Compared to WW walking, TM walking may thus require systematically elevated effort of gluteal muscles, in particular at slow walking speed.

Detailed spatial characterization of superficial hip muscle activation during walking: A multi-electrode surface EMG investigation of the gluteal region in healthy older adults.

PURPOSE: A multi-electrode array was used to generate spatially resolved Surface electromyography (SEMG) data of the hip muscles in healthy older adults. The cohort was meant to serve as an age-matched, normal control population for future surgical and rehabilitative studies in patients undergoing total hip arthroplasty, in view of the large, continuously increasing number of hip joint replacements. METHODS: Bilateral hip muscle SEMG activity, including tensor fasciae latae (TFL), gluteus medius (Gmed), and gluteus maximus (Gmax), was measured during locomotion on a walkway at self-selected slow, normal, and fast walking speeds (age-matched cohort of 29 females and 25 males). Eight equally-spaced, vertically oriented bipolar channels were applied on a horizontal line at mid-distance between iliac crest and greater trochanter (length 17.5 cm; named P1 to P8). Time-independent parameters (e.g., mean amplitude) were derived from the amplitude curves expressed as root mean square. RESULTS: The acquired SEMG data were not significantly influenced by gender (p = 0.202) or side (p = 0.313) and were therefore pooled. The most ventral to central electrode positions P1 to P5, representing TFL and ventral to central Gmed, showed the highest mean amplitude levels (averaged over the whole stride; 0.001 < p < 0.027 against P6 to P8; Bonferroni-adjusted paired t-test) at all walking speeds. Also, the respective curves showed two distinct amplitude peaks (representing load acceptance and hip stabilization during mid-stance), with a continuous increase of the first peak from P1 to P4 (most pronounced at fast speed) and the second peak from P1 to P3. Independently of the underlying individual muscles, both peaks displayed a continuous time shift from the most dorsal P8 to the most ventral P1 position, with the peaks for the ventral positions occurring at later time points during the normalized stride. CONCLUSIONS: The continuously changing activation patterns of the superficial muscles in the gluteal region during walking may reflect function-driven, finely tuned coordination patterns of neighboring muscles and muscle segments, rather than independent activation of anatomically defined muscles. This may be important for the definition of specific target parameters for the improvement and/or normalization of muscle function during training and post-injury rehabilitation.

Correction of static axial alignment in children with knee varus or valgus deformities through guided growth: Does it also correct dynamic frontal plane moments during walking?

Malaligned knees are predisposed to the development and progression of unicompartmental degenerations because of the excessive load placed on one side of the knee. Therefore, guided growth in skeletally immature patients is recommended. Indication for correction of varus/valgus deformities are based on static weight bearing radiographs. However, the dynamic knee abduction moment during walking showed only a weak correlation to malalignment determined by static radiographs. Therefore, the aim of the study was to measure the effects of guided growth on the normalization of frontal plane knee joint moments during walking. 15 legs of 8 patients (11-15 years) with idiopathic axial varus or valgus malalignment were analyzed. 16 typically developed peers served as controls. Instrumented gait analysis and clinical assessment were performed the day before implantation and explantation of eight-plates. Correlation between static mechanical tibiofemoral axis angle (MAA) and dynamic frontal plane knee joint moments and their change by guided growth were performed. The changes in dynamic knee moment in the frontal plane following guided growth showed high and significant correlation to the changes in static MAA (R=0.97, p<0.001). Contrary to the correlation of the changes, there was no correlation between static and dynamic measures in both sessions. In consequence two patients that had a natural knee moment before treatment showed a more pathological one after treatment. In conclusion, the changes in the dynamic load situation during walking can be predicted from the changes in static alignment. If pre-surgical gait analysis reveals a natural load situation, despite a static varus or valgus deformity, the intervention must be critically discussed.

Pair-rule patterning in the honeybee Apis mellifera: Expression of even-skipped combines traits known from beetles and fruitfly.

We have studied the binding pattern of antibody mAB 2B8 directed against even-skipped orthologous proteins (EVE) in honeybee embryos. Primary and secondary EVE stripes form in roughly anterior-to-posterior succession; there are 8 primary and 16 secondary stripes. The most posterior primary stripes appear only after the onset of gastrulation. The secondary stripes form by a splitting of primary stripes; they demarcate the parasegmental pattern. While these findings resemble EVE expression in long-germ beetles, the honeybee differs from both beetles and dipterans by two transient pair-rule traits in the parasegmental EVE pattern: the secondary stripes in head and thorax alternate in strength, yet out of register with the Drosophila pattern, and over the whole pattern the odd-numbered stripes vanish earlier than their even-numbered counterparts. As in Drosophila, however, the strong EVE stripes coincide with the weak engrailed (EN) stripes. These findings are taken to indicate that (1) honeybee and beetles share a conserved mode of EVE stripe formation whilst Drosophila has diverged in this respect, (2) honeybee and Drosophila have diverged from the beetles in specific pair-rule traits during the parasegmental expression of both EVE and EN, and (3) some of these traits differ in the register of segment pairing and thus may reflect regulatory divergences at the pair-rule level between dipterans and the honeybee.

Ernst Haeckel's ontogenetic recapitulation: irritation and incentive from 1866 to our time.

Ernst Haeckel (1834-1919) must count among the most widely known biologists of his time. His monographs on radiolarian skeletons, sponges and medusae immediately became standard works, owing partly to lavish illustrations that later on culminated in his "Art Forms in Nature", which markedly influenced the "art nouveau" of the early 20th century. Haeckel's main impact, however, came from his numerous popular books that were crucial in transferring Darwin's ideas to continental Europe. Haeckel's main addition was his claim that ontogeny recapitulates phylogeny. It was partly founded on pre-Darwinian observations by J.F. Meckel and K.E. v. Baer who noticed that vertebrate embryos of different species resemble each other more strongly during early ontogenesis than later on. Wishing to illustrate this clearly, Haeckel clandestinely generalized some figures showing early embryos of animals and Man. This "fraud" provided ammunition for his many adversaries, most of whom felt provoked by his antireligious campaigns. The resulting controversies continued well into the 20th century but then subsided. Quite recently, however, they have flared up again, perhaps in connection with progress in molecular embryology that revealed an amazing evolutionary conservation of genes and their cooperation in signal transduction chains. The scientific publications that triggered this flare, and a selection of "Letters to the Editor" in both international science magazines and the German popular press, serve here to show that Haeckel's idea of recapitulation, while having proven its heuristic value, is still causing considerable irritation. This results from the widespread intuition that the marvels of ontogenesis and other biological phenomena must reflect some "intelligent design" rather than Darwinian evolution.

Coagulation and fibrinolysis during lung surgery:an experimental study.

OBJECTIVES: Postoperative thromboembolism is a serious complication, but the incidence following surgery for lung cancer appears to be much lower compared with other surgical specialties. The reason is unknown and one may speculate that the lungs are reservoirs of anticoagulants or fibrinolytic substances, which are released by manipulation of the lung parenchyma during surgery. METHODS: Standardized lung manipulation, single-lung ventilation and pneumonectomy were performed in 10 anaesthetized pigs. Baseline and serial postmanipulation intravenous and intra-arterial blood samples were analysed for a wide range of fibrinolytic substances as well as pro- and anticoagulant factors. RESULTS: We found a transient but significant decrease in activated partial thromboplastin time (aPTT) and plasminogen activator inhibitor following manipulation of the lungs. Pneumonectomy resulted in minor but significant decrease in antithrombin and a significant increase in aPTT. All other measured substances were virtually constant. CONCLUSIONS: A wide range of fibrinolytic and anticoagulant substances remained unchanged during experimental lung manipulation. Minor changes were transient and not considered clinically relevant. Future studies should be initiated in humans because we have no deeper understanding why thromboembolic complications are relatively uncommon after general thoracic surgery compared with other surgical specialties.

An American in Paris and the origins of the stereomicroscope.

A century ago, Horatio S. Greenough, an American living in Paris, persuaded the firm of Carl Zeiss to construct the first low-power stereomicroscope. Fitted by Zeiss with Porro prisms for image erection, this instrument became the ancestor of all stereoscopic dissection microscopes ("binoculars") now in use. On the basis of original documents and earlier publications, the present Centennial Essay traces both the history of the Greenough stereomicroscope and the scantily documented life of its inventor, including his probable descent from a family of famous New England sculptors. The aim is to provide some historical background for a truly timeless instrument that was fundamental to developmental biology, beginning almost from the days of its proclamation by Wilhelm Roux.

Pattern formation in longitudinal halves of leaf hopper eggs (Homoptera) and some remarks on the definition of "Embryonic regulation".

1. Eggs of the leaf hopperEuscelis plebejus F. were fragmented longitudinally by a novel technique during cleavage. 2. Irrespective of the plane of fragmentation, both fragments may produce germ bands which are either of perfect bilateral symmetry or contain at least some bilaterally symmetrical body segments. With one possible exception, no longitudinal half-patterns were formed. 3. Ventral and lateral egg halves produce either complete germ bands or anterior partial germ bands. The failure to produce posterior body segments is correlated to the absence of special posterior pole materials marked by the symbiont mass. 4. Dorsal egg halves may produce either complete germ bands or posterior partial germ bands. The loss of anterior segments may be due to lack of keimanlagen cells. 5. The results are discussed and compared to those from transverse fragmentations carried out earlier. 6. Aided by a formalized presentation of the results a dilemma of definition is pointed out which results from the common usage of defining embryonic regulation as the alternative of mosaic development and at the same time by reference to the normal result of development.

Aberrant oogenesis in the patterning mutant dicephalic of Drosophila melanogaster: time-lapse recordings and volumetry in vitro.

Drosophila females homozygous for the mutation dicephalic occasionally produce ovarian follicles with a nurse-cell cluster on each oocyte pole (dic follicles). Most dic follicles contain 15 nurse cells as in the normal follicle, but the total nurse-cell volume is larger in dic follicles; this is in keeping with the increase in DNA content recently described. However, the relative increase in oocyte volume during nurse-cell regression (from stage 10B onward) is not significantly larger in dic than in normal follicles. Time-lapse recordings in vitro show that, as a rule, both nurse cell clusters in a dic follicle export cytoplasm to the oocyte but nurse-cell regression remains incomplete at both poles and the persisting remnants of the nurse cells cause anomalies in chorion shape. The kinematics of cytoplasmic transfer are less aberrant at that oocyte pole which harbours the germinal vesicle. Possible links are discussed between these anomalies of oogenesis and the double-anterior embryonic patterns observed in the majority of developing dic eggs.

[Development of the malformation "double abdomen" in eggs of smittia parthenogenetica (dipt., chironomidae) partially irradiated by UV]. / Der Entwicklungsgang der Mißbildung "Doppelabdomen" im partiell UV-bestrahlten Ei von Smittia parthenogenetica (Dipt., Chironomidae).

UV-irradiation (maximum intensity at 253,7 nm) of the anterior 1/3 of the egg during nuclear migration stage frequently is followed by the formation of "double abdomens". The type of malformation shows a germ band in which head, thorax and anterior abdominal segments are replaced by a set of 6-8 posterior abdominal segments joined in mirror-image symmetry to the corresponding parts of the original abdomen. The course of development leading to this malformation is visibly aberrant from the blastoderm stage to germ band segmentation only. The two abdomens develop in strict synchrony provided their positions are symmetrical to the equatorial plane of the egg. Asymmetric positions are less frequent and delay temporarily the development of one or the other of the abdomens due to the lack of space. The results are compared to similar malformations in other insect eggs and possible interpretations are reviewed.

Evo-devo aspects of classical and molecular data in a historical perspective.

We discuss the interplay between evolution and development as reflected in data and concepts since about 1800. Darwin and his "continental apostle" Haeckel put the striking similarity between early vertebrate embryos in an evolutionary context. Haeckel's partly illicit generalizations discredited evolutionary thinking among early experimental embryologists who moreover noted riddles incompatible with contemporary concepts of homology and evolution. Relevant solutions were suggested by the more recent concept of ontogenetic networks that embody complex regulatory properties and genes with partly overlapping functions. Molecular data on development increasingly reveal evolutionary opportunism, for instance when a widespread signaling chain involved in primitive immune defense was apparently recruited later on for dorso-ventral axis determination in some evolutionarily advanced insect groups. Recently, Rickettsia-related bacteria colonizing many arthropod species were found to "manipulate" the first steps of host development to the advantage of their own propagation, but by ways that could also promote host speciation. Molecular genetics can now document evolutionary steps in ontogenetic networks. In the fruit fly for instance, the novel bicoid gene has superseded a crucial patterning function within a pre-existing network--a case of "molecular caenogenesis." The expression patterns of conserved genes that antagonistically determine dorso-ventral polarity support a literal revolution envisioned almost 200 years ago. This is the dorso-ventral inversion of the body plan in some metazoans--ascribed then to the Articulata, now to the Chordata. The final section posits that the opportunistic character of evolutionary innovations is detrimental to parsimony in development.

The spatial arrangement of germ line cells in ovarian follicles of the mutantdicephalic inDrosophila melanogaster.

The pattern of intercellular connections between germ line cells has been studied in follicles of the mutantdicephalic (dic), which possess nurse cell clusters at both poles. Staining of follicles with a fluorescent rhodamine conjugate of phalloidin reveals ring canals and cell membranes and thus allows us to reconstruct the spatial organization of the follicle. Each germ line cell can be identified by the pattern of cell-cell connections which reflect the mitotic history of individual cells in the 16-cell cluster. The results indicate that in both wild-type anddicephalic cystocyte clusters one of the two cells with four ring canals normally becomes the pro-oocyte. However, in some follicles (dicephalic and wild-type) oocytes were found with fewer or more than four ring canals. Indic follicles, one or several nurse cells may become disconnected from the other cells during oocyte growth at stage 9-10. Such disconnected cells cannot later on empty their cytoplasm into the oocyte. This, in turn, might be of consequence for the determination of axial polarity of the embryo.

A dicephalic monster embryo ofDrosophila melanogaster.

A monster embryo with a head on each end developed from an abnormalDrosophila egg produced by a M(1)oSp/Binsn female. The embryo probably represents a longitudinal mirror duplication of head and all 3 thoracic segments. The significance of this type of pattern anomaly for embryonic pattern specification inDrosophila is discussed.