Predictive Simulations of Musculoskeletal Function and Jumping Performance in a Generalized Bird.

Jumping is a common, but demanding, behavior that many animals employ during everyday activity. In contrast to jump-specialists such as anurans and some primates, jumping biomechanics and the factors that influence performance remains little studied for generalized species that lack marked adaptations for jumping. Computational biomechanical modeling approaches offer a way of addressing this in a rigorous, mechanistic fashion. Here, optimal control theory and musculoskeletal modeling are integrated to generate predictive simulations of maximal height jumping in a small ground-dwelling bird, a tinamou. A three-dimensional musculoskeletal model with 36 actuators per leg is used, and direct collocation is employed to formulate a rapidly solvable optimal control problem involving both liftoff and landing phases. The resulting simulation raises the whole-body center of mass to over double its standing height, and key aspects of the simulated behavior qualitatively replicate empirical observations for other jumping birds. However, quantitative performance is lower, with reduced ground forces, jump heights, and muscle-tendon power. A pronounced countermovement maneuver is used during launch. The use of a countermovement is demonstrated to be critical to the achievement of greater jump heights, and this phenomenon may only need to exploit physical principles alone to be successful; amplification of muscle performance may not necessarily be a proximate reason for the use of this maneuver. Increasing muscle strength or contractile velocity above nominal values greatly improves jump performance, and interestingly has the greatest effect on more distal limb extensor muscles (i.e., those of the ankle), suggesting that the distal limb may be a critical link for jumping behavior. These results warrant a re-evaluation of previous inferences of jumping ability in some extinct species with foreshortened distal limb segments, such as dromaeosaurid dinosaurs. Simulations prédictives de la fonction musculo-squelettique et des performances de saut chez un oiseau généralisé Sauter est un comportement commun, mais exigeant, que de nombreux animaux utilisent au cours de leurs activités quotidiennes. Contrairement aux spécialistes du saut tels que les anoures et certains primates, la biomécanique du saut et les facteurs qui influencent la performance restent peu étudiés pour les espèces généralisées qui n'ont pas d'adaptations marquées pour le saut. Les approches de modélisation biomécanique computationnelle offrent un moyen d'aborder cette question de manière rigoureuse et mécaniste. Ici, la théorie du contrôle optimal et la modélisation musculo-squelettique sont intégrées pour générer des simulations prédictives du saut en hauteur maximal chez un petit oiseau terrestre, le tinamou. Un modèle musculo-squelettique tridimensionnel avec 36 actionneurs par patte est utilisé, et une méthode numérique nommée "direct collocation" est employée pour formuler un problème de contrôle optimal rapidement résoluble impliquant les phases de décollage et d'atterrissage. La simulation qui en résulte élève le centre de masse du corps entier à plus du double de sa hauteur debout, et les aspects clés du comportement simulé reproduisent qualitativement les observations empiriques d'autres oiseaux sauteurs. Cependant, les performances quantitatives sont moindres, avec une réduction des forces au sol, des hauteurs de saut et de la puissance musculo-tendineuse. Une manœuvre de contre-mouvement prononcée est utilisée pendant le lancement. Il a été démontré que l'utilisation d'un contre-mouvement est essentielle à l'obtention de hauteurs de saut plus importantes, et il se peut que ce phénomène doive exploiter uniquement des principes physiques pour réussir; l'amplification de la performance musculaire n'est pas nécessairement une raison immédiate de l'utilisation de cette manœuvre. L'augmentation de la force musculaire ou de la vitesse de contraction au-dessus des valeurs nominales améliore grandement la performance de saut et, fait intéressant, a le plus grand effet sur les muscles extenseurs des membres plus distaux (c'est-à-dire ceux de la cheville), ce qui suggère que le membre distal peut être un lien critique pour le comportement de saut. Ces résultats justifient une réévaluation des déductions précédentes de la capacité de sauter chez certaines espèces éteintes avec des segments de membres distaux raccourcis, comme les dinosaures droméosauridés. Voorspellende simulaties van musculoskeletale functie en springprestaties bij een gegeneraliseerde vogel Springen is een veel voorkomend, maar veeleisend, gedrag dat veel dieren toepassen tijdens hun dagelijkse bezigheden. In tegenstelling tot de springspecialisten zoals de anura en sommige primaten, is de biomechanica van het springen en de factoren die de prestaties beïnvloeden nog weinig bestudeerd voor algemene soorten die geen uitgesproken adaptaties voor het springen hebben. Computationele biomechanische modelbenaderingen bieden een manier om dit op een rigoureuze, mechanistische manier aan te pakken. Hier worden optimale controle theorie en musculoskeletale modellering geïntegreerd om voorspellende simulaties te genereren van maximale hoogtesprong bij een kleine grondbewonende vogel, een tinamou. Een driedimensionaal musculoskeletaal model met 36 actuatoren per poot wordt gebruikt, en directe collocatie wordt toegepast om een snel oplosbaar optimaal controleprobleem te formuleren dat zowel de opstijg-als de landingsfase omvat. De resulterende simulatie verhoogt het lichaamszwaartepunt tot meer dan het dubbele van de stahoogte, en belangrijke aspecten van het gesimuleerde gedrag komen kwalitatief overeen met empirische waarnemingen voor andere springende vogels. De kwantitatieve prestaties zijn echter minder, met verminderde grondkrachten, spronghoogtes en spierpeeskracht. Tijdens de lancering wordt een uitgesproken tegenbewegingsmanoeuvre gebruikt. Aangetoond is dat het gebruik van een tegenbeweging van cruciaal belang is voor het bereiken van grotere spronghoogten, en dit fenomeen hoeft alleen op fysische principes te berusten om succesvol te zijn; versterking van de spierprestaties hoeft niet noodzakelijk een proximate reden te zijn voor het gebruik van deze manoeuvre. Het verhogen van de spierkracht of van de contractiesnelheid boven de nominale waarden verbetert de sprongprestatie aanzienlijk, en heeft interessant genoeg het grootste effect op de meer distale extensoren van de ledematen (d.w.z. die van de enkel), wat suggereert dat de distale ledematen een kritieke schakel kunnen zijn voor het springgedrag. Deze resultaten rechtvaardigen een herevaluatie van eerdere conclusies over springvermogen bij sommige uitgestorven soorten met voorgekorte distale ledematen, zoals dromaeosauride dinosauriërs. Prädiktive Simulationen der muskuloskelettalen Funktion und Sprungleistung bei einem generalisierten Vogel Springen ist ein übliches jedoch anstrengendes Verhalten, das viele Tiere bei ihren täglichen Aktivitäten einsetzen. Im Gegensatz zu Springspezialisten, wie Fröschen und einigen Primaten, sind bei allgemeinen Arten, welche keine ausgeprägten Anpassung für Sprungverhalten aufweisen, die Biomechanik beim Springen und die Faktoren, welche die Leistungsfähigkeit beeinflussen, noch wenig untersucht. Computergestützte biomechanische Modellierungsverfahren bieten hier eine Möglichkeit, dies in einer gründlichen, mechanistischen Weise anzugehen. In dieser Arbeit werden die optimale Steuerungstheorie und Muskel-Skelett-Modellierung zusammen eingesetzt, um die maximale Sprunghöhe eines kleinen bodenlebenden Vogels, eines Perlsteisshuhns, zu simulieren und zu prognostizieren. Es wird ein dreidimensionales Muskel-Skelett-Modell mit 36 Aktuatoren pro Bein verwendet, und durch direkte Kollokation wird ein schnell lösbares optimales Steuerungsproblem formuliert, das sowohl die Abstoss- als auch die Landephase umfasst. Die daraus folgende Simulation bringt den Ganzkörperschwerpunkt auf mehr als das Doppelte seiner Standhöhe und entscheidende Aspekte des simulierten Verhaltens entsprechen qualitativ empirischen Beobachtungen für andere springende Vögel. Allerdings ist die quantitative Leistungsfähigkeit geringer, mit reduzierten Bodenkräften, Sprunghöhen und Muskel-Sehnen-Kräften. Beim Abstossen wird ein ausgeprägtes Gegenbewegungsmanöver durchgeführt. Die Durchführung einer Gegenbewegung ist nachweislich entscheidend für das Erreichen grösserer Sprunghöhen, wobei dieses Phänomen möglicherweise nur physikalische Prinzipien auszuschöpfen braucht, um erfolgreich zu sein. Die Verstärkung der Muskelleistung ist daher möglicherweise nicht zwingend ein unmittelbarer Grund für die Verwendung dieses Manövers. Eine Erhöhung der Muskelkraft oder der Kontraktionsgeschwindigkeit über die Nominalwerte hinaus führt zu einer erheblichen Zunahme der Sprungleistung und hat interessanterweise den grössten Effekt bei den weiter distal gelegenen Streckmuskeln der Beine (d.h. bei denjenigen des Sprunggelenks), was darauf hindeutet, dass die distale Gliedmasse ein entscheidendes Element für das Sprungverhalten sein könnte. Diese Ergebnisse geben Anlass zur Überprüfung früherer Schlussfolgerungen hinsichtlich der Sprungfähigkeit einiger ausgestorbener Arten mit verkürzten distalen Gliedmassen, wie beispielsweise bei dromaeosauriden Dinosauriern.

Effect of irrigation at critical stages on the phenology of flowering and fruiting of the cactus Opuntia spp / Efeito da irrigação em estágios críticos sobre a fenologia de floração e frutificação do cacto Opuntia spp

Abstract This paper briefly reports some effects of irrigations at two critical periods on the phenology of three varieties of cactus pear cultivated in Agadir area: the spineless varieties 'Aissa' and 'Moussa' and the thorny one 'Achefri'. In the first year experiments (2010-2011) treatments of irrigation used were: (T1) 0 mm, (T2) 30 mm during flowering and 30 mm during fruit enlargement and (T3) 30 mm only during fruit enlargement. In the 2nd year experiments, irrigation treatments were: (T1) 0 mm, (T2) 60 mm during flowering and 60 mm during fruit enlargement and (T3) 60 mm only during fruit enlargement. Treatments of irrigation were applied between mid-April and mid-June in the 1st year experiments and in February and May in the 2nd year experiments. Results of the first year experiments showed that the emission of buds was higher in the thorny variety than in the spineless ones (more than 6 emitted buds/cladode vs less than 4.5 in the spineless varieties). In the second year, irrigation increased the emission of buds in the three varieties (more than 7 emitted buds/cladode for each T2 and T3 of all varieties vs not more than 5 for T1) and the duration of the flowering phase of these varieties. However, irrigation did not modify the proportions of fruits reaching commercial maturity during the early or the late period of maturation.

Resumo Este artigo relatou brevemente alguns efeitos das irrigações em dois períodos críticos na fenologia de três variedades de Opuntia cultivadas na área de Agadir: as variedades sem espinhos 'Aissa' e 'Moussa' e um espinhoso 'Achefri'. No primeiro ano (2010-2011) os seguintes tratamentos do experimento da irrigação foram usados: (T1) 0 milímetros, (T2) 30 milímetros durante a florescência e 30 milímetros durante a ampliação do fruto e (T3) 30 milímetros somente durante a ampliação do fruto. No segundo ano das experiências, os tratamentos da irrigação foram: (T1, 0 milímetros), (T2) 60 milímetros durante a florescência e 60 milímetros durante a ampliação do fruto e (T3) 60 milímetros somente durante a ampliação do fruto. Os tratamentos da irrigação foram aplicados entre a metade de abril e metade de junho nas experiências do primeira ano e em fevereiro e maio nas experiências do segundo ano. Os resultados das experiências do primeiro ano mostraram que a emissão de brotos foi maior na variedade espinhosa do que no rufia (mais de 6 emitida gomos/cladode vs inferior a 4,5 no rufia variedades). No segundo ano, a irrigação aumentou a emissão dos botões nas três variedades (mais de 7 botões emitidos/cladÓdio para cada T2 e T3 de todas as variedades não mais que 5 para o T1) e a duração da fase de florescência destas variedades. Contudo, a irrigação não alterou as proporções de frutos que alcançaram a maturidade comercial durante o período adiantado ou atrasado de maturação.

Effect of irrigation at critical stages on the phenology of flowering and fruiting of the cactus Opuntia spp.

This paper briefly reports some effects of irrigations at two critical periods on the phenology of three varieties of cactus pear cultivated in Agadir area: the spineless varieties 'Aissa' and 'Moussa' and the thorny one 'Achefri'. In the first year experiments (2010-2011) treatments of irrigation used were: (T1) 0 mm, (T2) 30 mm during flowering and 30 mm during fruit enlargement and (T3) 30 mm only during fruit enlargement. In the 2nd year experiments, irrigation treatments were: (T1) 0 mm, (T2) 60 mm during flowering and 60 mm during fruit enlargement and (T3) 60 mm only during fruit enlargement. Treatments of irrigation were applied between mid-April and mid-June in the 1st year experiments and in February and May in the 2nd year experiments. Results of the first year experiments showed that the emission of buds was higher in the thorny variety than in the spineless ones (more than 6 emitted buds/cladode vs less than 4.5 in the spineless varieties). In the second year, irrigation increased the emission of buds in the three varieties (more than 7 emitted buds/cladode for each T2 and T3 of all varieties vs not more than 5 for T1) and the duration of the flowering phase of these varieties. However, irrigation did not modify the proportions of fruits reaching commercial maturity during the early or the late period of maturation.

Effect of irrigation at critical stages on the phenology of flowering and fruiting of the cactus Opuntia spp

Abstract This paper briefly reports some effects of irrigations at two critical periods on the phenology of three varieties of cactus pear cultivated in Agadir area: the spineless varieties Aissa and Moussa and the thorny one Achefri. In the first year experiments (2010-2011) treatments of irrigation used were: (T1) 0 mm, (T2) 30 mm during flowering and 30 mm during fruit enlargement and (T3) 30 mm only during fruit enlargement. In the 2nd year experiments, irrigation treatments were: (T1) 0 mm, (T2) 60 mm during flowering and 60 mm during fruit enlargement and (T3) 60 mm only during fruit enlargement. Treatments of irrigation were applied between mid-April and mid-June in the 1st year experiments and in February and May in the 2nd year experiments. Results of the first year experiments showed that the emission of buds was higher in the thorny variety than in the spineless ones (more than 6 emitted buds/cladode vs less than 4.5 in the spineless varieties). In the second year, irrigation increased the emission of buds in the three varieties (more than 7 emitted buds/cladode for each T2 and T3 of all varieties vs not more than 5 for T1) and the duration of the flowering phase of these varieties. However, irrigation did not modify the proportions of fruits reaching commercial maturity during the early or the late period of maturation.

Resumo Este artigo relatou brevemente alguns efeitos das irrigações em dois períodos críticos na fenologia de três variedades de Opuntia cultivadas na área de Agadir: as variedades sem espinhos 'Aissa' e 'Moussa' e um espinhoso 'Achefri'. No primeiro ano (2010-2011) os seguintes tratamentos do experimento da irrigação foram usados: (T1) 0 milímetros, (T2) 30 milímetros durante a florescência e 30 milímetros durante a ampliação do fruto e (T3) 30 milímetros somente durante a ampliação do fruto. No segundo ano das experiências, os tratamentos da irrigação foram: (T1, 0 milímetros), (T2) 60 milímetros durante a florescência e 60 milímetros durante a ampliação do fruto e (T3) 60 milímetros somente durante a ampliação do fruto. Os tratamentos da irrigação foram aplicados entre a metade de abril e metade de junho nas experiências do primeira ano e em fevereiro e maio nas experiências do segundo ano. Os resultados das experiências do primeiro ano mostraram que a emissão de brotos foi maior na variedade espinhosa do que no rufia (mais de 6 emitida gomos/cladode vs inferior a 4,5 no rufia variedades). No segundo ano, a irrigação aumentou a emissão dos botões nas três variedades (mais de 7 botões emitidos/cladÓdio para cada T2 e T3 de todas as variedades não mais que 5 para o T1) e a duração da fase de florescência destas variedades. Contudo, a irrigação não alterou as proporções de frutos que alcançaram a maturidade comercial durante o período adiantado ou atrasado de maturação.