in Crested Ducks (Anas platyrhynchos f. d.)
Transcription
in Crested Ducks (Anas platyrhynchos f. d.)
Arch.Geflügelk., 74 (3). S. 203–209, 2010, ISSN 0003-9098. © Verlag Eugen Ulmer, Stuttgart Brain alterations, their impact on behavior and breeding strategy in Crested Ducks (Anas platyrhynchos f. d.) Hirnveränderungen, ihr Einfluss auf das Verhalten und Zuchtmanagement bei haubentragenden Hausenten (Anas platyrhynchos f. d.) Julia Mehlhorn and G. Rehkämper PhD Award 2009 of German Branch of WPSA Introduction Domestication could be seen as a selective process associated with a lot of alterations and greater variability in a lot of signs in domestic animals if compared to their wild ancestors (PRICE, 1999). During the course of domestication a lot of breeds have been developed which show alterations in e.g. body size, coloring, habitat or behavior. Also alterations can be found in brain size and brain composition (KRUSKA, 1980; EBINGER, 1995), not only in comparison to wild animals but also between different breeds (REHKÄMPER et al., 2003, 2008). One example for this is the Crested Duck (CR hereafter, Figure 1), which is characterized by a feather crest. The existence of a crest can be associated with changes in skull and brain morphology. In crested chickens the frontal bones form a protuberance, into which the telencephalon is displaced and show peculiarities in size and composition (FRAHM and REHKÄMPER, 1998). In contrast, the crest in CR is located in the parietal part of the skull underlain by a cushion of fat and connective tissue. Additionally, many CR bear a fat body inside the skull that could vary in size and position (Figure 2). Depending on its size and position in relation to the brain, a negative effect on brain function cannot not be excluded and in fact, often coordination of locomotion is poor as demonstrated by a tottering walk, and some animals are even unable to right themselves up after fallen on their back (BRINKMEIER, 1999; BARTELS and KUMMERFELD, 2001; BARTELS et al., 2001; FRAHM et al., 2001; CNOTKA et al., 2006, 2007, 2008). Hereditary aspects and high pre- and postnatal mortalities were discussed by BARTELS and KUMMERFELD (2001) and BARTELS et al. (2001). The German government has asked experts to work out some guidelines for the breeding of fancy poultry (“Gutachten zur Auslegung von § 11b des Tierschutzgesetzes [Verbot von Qualzüchtungen]”). These guidelines discuss crested ducks critically and propose to consider prohibiting the breeding of CR with regards to the German law preventing cruelty against animals. In this study an analysis of the observed problems, their possible origin and of the brains of CR was done. Additionally a behavioral test and a breeding strategy had to be developed that would allow a breeder to discover animals Working Group Behavior and Brain, C.&O. Vogt Institute of Brain Research, University of Düsseldorf, Germany Arch.Geflügelk. 3/2010 with large fat bodies even if the undesirable defect is not obvious and behavior in daily life looks quite normal. The goal is to eliminate detrimental traits from their breeding populations, assuming that the condition is heritable (LANCASTER, 1990). It is proven that the existence of a crest and its size is not a reliable predictor of the fat body or behavioral deficits (KRAUTWALD, 1910; BRINKMEIER, 1999; BARTELS and KUMMERFELD, 2001; FRAHM et al., 2001, 2005) and thus, other phenotypic measures must be identified that are associated with this trait. A good breeding management should solve the existing problems rather than eliminating this old and traditional breed completely. The present paper draws attention to a behavioral test that indicates deficits in coordination of locomotion. To prove the appropriateness of this test, individuals preselected by the test were used in a special breeding program. Number of offspring, coordination of locomotion and morphology of skull and brain were recorded. In parallel the brain size and brain composition was analyzed in detail in this breed and it was checked whether individuals identified as malfunctional have a more or less prominent fat body inside their skulls. Material and Methods Behavioral test In total, 26 adult CR (11 drakes, 15 ducks) from our own stock were used for behavioral test and brain analysis. The behavioral test should check the motor abilities and should challenge the system of motor coordination of CR. The idea for the used ‘righting test’ test was based on the fact that ducks with serious problems in locomotion coordination often fall down on their back and then are unable to right themselves and stand up. Among the 26 ducks investigated, ten individuals showed motor incoordination such as staggering during moving or tumbling down at least one time in their lives. These ducks exhibited the symptoms in different manners but appeared quite normal in their daily lives. The deficits became apparent during stressful situations like collecting or handling of the ducks. After such treatment, many of them recovered quickly and any deficits were inconspicuous again in daily life. The other 16 ducks had no detectable deficits in motor coordination and were completely inconspicuous in their locomotion behavior. So, the following ‘righting test’ was employed on two distinguishable groups of ducks, “affected” CR (n = 10) and “normal” CR (n = 16). For the test, all CR were placed on their backs and the required time needed to stand up on 204 Mehlhorn und Rehkämper: Brain alterations and behaviour in Crested Ducks Figure 1. Portrait of a Crested Duck (CR) Kopfstudie einer Landente mit Haube Figure 2. Dorsal view of a Crested Duck brain with incorporated middle sized fat body (*) Landentengehirn mit einem Fettkörper mittlerer Größe (*), von dorsal gesehen both feet was recorded. This procedure was repeated 13 times per animal over a period of three months. The average time was calculated for each individual. Differences between the 2 examined groups for average righting time was calculated and evaluated using the Mann-Whitney Rank Sum Test. body sizes to correct for the influence of body size. This method involves the calculation of a regression line slope that expresses the brain (or brain component or fat body) size/body size relationship. To test for differences in brain structure volumes between CR and pooled reference breeds allometric size indices [(brain component size/expected brain component size) * 100] were calculated. The indices represented the distance of individual data points from the regression line. The expected brain (or brain component size) is the value on the regression line that corresponds to a given individual body weight. A detailed description of these methods for preparation, measurement and calculation is given in literature (STEPHAN et al., 1988; FRAHM and REHKÄMPER, 1998; MPOTSARIS and REHKÄMPER, 2006; CNOTKA et al., 2007). Differences between the 2 examined groups for fat body volume were calculated and evaluated using the MannWhitney Rank Sum Test. Differences between the size indices of CR and the reference group were evaluated using Student’s t-Test. Additionally correlation analyses between fat body volume and brain size or brain component size were calculated using Pearson product moment correlation. Brain analysis The investigation of the brain anatomy was based on 26 CR brains from the tested animals. For comparison 26 brains from three other domestic duck breeds, called Pommeranian Ducks (Po, n = 10, 5 drakes/5 ducks), Abacot Ranger Ducks (Ar, n = 6, 3 drakes/3 ducks) and “Hochbrutflugenten” (Hb, n = 10, 5 drakes/5 ducks) were used. These breeds represent the body size variability in domestic ducks which will be important for brain analyses with allometric methods. All these breeds were uncrested and do not show, as far as known, any peculiarities like those observed in CR. All specimens of ducks were obtained from organized breeders and were at least one year old. The animals were euthanized with an overdose of pent barbiturate. After cardiac arrest was confirmed ducks were perfused with physiological saline solution to wash out the blood, followed by Bodian’s fluid to fixate the brain. After this, the brains were carefully dissected, sectioned and stained for cell bodies. Body weight, total brain volume, volume of the fat body and the volume of 14 other areas of the brain (hyperpallium apicale, hyperpallium densocellulare, mesopallium, nidopallium, striatopallidal complex, septum, hippocampus, area prepiriformis, bulbus olfactorius, tegmentum, cerebellum, tectum, tractus opticus, diencephalon) were determined. For volume measurements the contours of brain structures were drawn using a digital pen and a camera lucida. For calculation of the fresh volume, the resulting values were multiplied by the section thickness, the distance between the sections and the individual conversion factor of shrinkage (volume fresh brain/sum of serial section volumes). Net brain volume was calculated as the sum of the single brain components. In contrast to total brain volume, the net brain volume does not include the volume of leptomeninges, ventricles, choroids plexus and remains of brain nerves. Allometric methods were used for comparison of volumes of brain structures in different breeds with different Breeding In total, three years of breeding have been completed. In the first year three breeding couples were chosen from our own stock without any special selection. In the following two years the breeding couples (five in the 2nd year and three in the 3rd year) were preselected by the ‘righting test’. Only individuals with the shortest average righting times were chosen for breeding. All breeding couples were housed in a separate aviary with their own swimming facility. The eggs were collected every day and hatched by a common incubator. The ducklings were reared all together in a separate aviary with good observation conditions. Eggs that did not hatch were opened three days after the end of hatching time and the habitus and skull of extant embryos were investigated. Thus, fertilization rate, hatching rate, number of unhatched ducklings with skull malformations (e.g. perforations of the skull) and number of (hatched) ducklings with behavioral deficits of these breeding couples were investigated. Hatched ducks were checked for skull deformations after growing up as well. Arch.Geflügelk. 3/2010 Mehlhorn und Rehkämper: Brain alterations and behaviour in Crested Ducks Differences in the investigated parameters between the three breeding years were calculated using Chi2-Test. All research was performed in accordance with the official German regulations for research on animals. Results Behavioral test The ‘righting test’ was employed on two distinguishable groups of ducks (see above): the 10 individuals with several observed deficits in coordination of locomotion and the 16 individuals without any behavioral deficits. In contrast to the “normal” ducks, the “affected” CR performed poorly. On average they required 13.4 s ± 6.93 with a range from 1.0 to 62.6 s for the ‘righting test’. The “normal” CR required 1.4 s ± 0.20 s and ranged from 0.5 to 2.9 s. The average times of the two groups differed significantly (t = 2.21; DF = 24; p = 0.037). Brain analysis On average the body weight of all affected CR investigated amounted to 2,301 g ± 58.4 and ranged from 1,725 to 2,960 g. The corresponding data on brain size are 6,855 mm3 ± 209 with a range from 5,111 mm3 to 9,547 mm3. Normal CR ducks show an average body weight of 2,300 g ± 81.7 (range from 1,825 to 2,960 g) and an average brain size of 6,659 mm3 ± 225 (range from 5,111 to 8,060 mm3). Two (normal) CR ducks from the examined duck population did not show a fat body. The volumes of the fat bodies of the other 24 CR ducks varied from 19 mm3 to 3,891 mm3 and from 0.3% to 41% of total brain volume, respectively. The fat body size of the two groups (consisting of 10 “affected” and 16 “normal” ducks) differed significantly. Ducks with deficits in coordination of locomotion showed a significantly larger fat body (1,512 mm3 ± 371) than ducks without such problems (637 mm3 ± 138; 2-tailed t-test, t = 2.583, DF = 24, p = 0.016). Most frequently mid- 205 dle-sized fat bodies were positioned in the tentorium cerebelli (Figure 2). Additionally, the fat bodies often extended to rostral between the two hemispheres (Figure 3). There was no significant correlation between the fat body size and the latency to right (r = 0.278, p = 0.169). The allometric comparison of the brain and brain structure volume in relationship to the body weight of all ducks is represented in Table 1. CR individuals (n = 26) exhibit a significantly larger total brain volume in comparison to the pooled reference ducks (n = 26, t = 2.651, DF = 50, p = 0.011). The total brain volume included, if present, the volume of the fat body. But, as can be seen in Table 1 and Figure 4, net brain volume of CR was significantly smaller in an allometric comparison to the reference breeds (t = –2.325, DF = 50, p = 0.024). Among the 14 brain structures delineated, in CR 10 were not reduced compared to reference breeds. A relative reduction of cerebellum, tegmentum, apical hyperpallium and olfactory bulb was observed in comparison to the reference group (cerebellum: t = –2.882, DF = 50, p = 0.006; tegmentum: t = –2.567, DF = 50, p = 0.013; apical hyperpallium: t = –2.056, DF = 50, p = 0.045; olfactory bulb: t = –4.392, DF = 48, p < 0.001). The correlation analysis revealed an expected clear positive correlation between the fat body volume and the total brain volume of CR (r = 0.908, n = 26, p < 0.001). There were no significant correlations between the fat body size and the reduced net brain (net brain without fat, r = –0.271, p = 0.18) or the reduced structures of cerebellum (r = –0.142, p = 0.49), tegmentum (r = –0.0147, p = 0.949), bulbus olfactorius (r = –0.0759, p = 0.725) or hyperpallium apicale (r = –0.174, p = 0.394). Breeding Breeding results are presented in Table 2. The average hatching rate of the (randomly chosen) breeding couples of the first year (64%) was significantly different to that of the next two years (86%, 1st year vs. 2nd year: Chi2 = 8.97, z = 2.995, p = 0.003; 1st year vs. 3rd year: Chi2 = 6.248, z = 2.5, p = 0.012). The number of unhatched ducklings 10000 Net brain volume [mm3] 9000 8000 Crested Ducks "Hochbrutflugenten" Pommeranian Ducks Abacot Ranger Ducks 7000 6000 5000 a = 0,167 b = 3,183 4000 1000 2000 3000 Body weight [g] Figure 3. Coronal section through the brain of a Crested Duck with an intracranial fat body Frontalschnitt durch das Gehirn einer Landente mit intrakranialem Fettkörper Arch.Geflügelk. 3/2010 Figure 4. Exemplary double logarithmic plot of net brain volume vs. body weight. Individual data points, averages and standard deviations are given for four duck breeds (explanations in the text) Doppelt logarithmierter Graph als Beispiel für die Beziehung zwischen Nettohirnvolumen und Körpergewicht. Dargestellt sind Einzelwerte, Mittelwerte und Standardabweichungen für vier Hausentenrassen (Erklärungen im Text) 206 Mehlhorn und Rehkämper: Brain alterations and behaviour in Crested Ducks Table 1. Volumes of brain structures (mm3) and allometric size indices in 4 breeds of domestic ducks (CR = Crested Ducks, Hb = “Hochbrutflugenten”, Po = Pommeranian Ducks, Ar = Abacot Ranger Ducks (means ± S.D.; * p ≤ 0.05, ** p ≤ 0.01, *** p ≤ 0.001) Hirnstukturvolumina (mm3) und allometrische Größenindices von 4 Hausentenrassen (CR = Landenten mit und ohne Haube, Hb = Hochbrutflugenten, Po = Pommernenten, Ar = Streicherenten; Mittelwerte ± SD; * p ≤ 0,05, ** p ≤ 0,01, *** p ≤ 0,001) Crested Ducks Structure (n = 26) Volume “HochbrutPommeranian flugenten” Ducks (n = 10) (n = 10) ± 209 4913 ± 89.8 4779 ± 65.7 ± 65.9 Hyperpallium apicale 520 Hyperpallium densocellulare 159 Hyperpallium ventrale 547 Nidopallium 1643 Striatopallidal complex 485 Hippocampus 78.8 Septum 35.9 Prepiriformis 16.4 Bulbus olfactorius 26.8 ± ± ± ± ± ± ± ± ± 14.6 3.68 10.7 38.3 14.2 3.14 1.23 0.56 0.59 442 134 479 1520 420 76.8 30.4 12.6 25.4 ± 7.11 520 ± 2.13 157 ± 8.10 547 ± 22.9 1623 ± 5.73 485 ± 1.70 84.5 ± 0.63 38.0 ± 0.37 15.7 ± 0.61 33.6 Telencephalon Diencephalon Tractus opticus Tectum Tegmentum Cerebellum ± 76.2 ± 6.05 ± 2.12 ± 4.29 ± 10.2 ± 11.5 3140 241 63.1 213 633 490 ± 39.4 3503 ± 4.52 310 ± 1.93 80.5 ± 3.98 254 ± 11.3 759 ± 14.3 682 Total brain volume Net brain volume 6846 5405 3511 293 70.5 232 693 606 ± 132 ± 123 5814 5589 Abacot Ranger Ducks (n = 6) Indices Hb + Po + Ar Crested Ducks (n = 26) (n = 26) ± 112 ± 153 98.0 ± 1.32 103 ± 1.53 ± 9.40 677 ± 7.23 171 ± 20.4 625 ± 36.0 1967 ± 12.5 534 ± 1.85 72.5 ± 0.76 36.0 ± 0.48 15.5 ± 1.03 33.7 ± 8.86 ± 8.82 ± 17.7 ± 42.3 ± 7.69 ± 3.69 ± 1.96 ± 1.55 ± 2.44 105 102 101 103 101 101 102 99.7 109 ± ± ± ± ± ± ± ± ± 2.91 2.45 2.22 2.09 1.69 1.94 1.69 2.68 3.14 96.9 101 97.9 98.4 99.9 101 99.2 103 92.5 ± 2.66* ± 2.34 ± 1.92 ± 2.30 ± 3.03 ± 4.27 ± 3.38 ± 3.62 ± 2.12*** ± 78.0 ± 6.65 ± 2.79 ± 8.46 ± 22.4 ± 14.2 ± ± ± ± ± ± 103 102 104 102 103 105 ± ± ± ± ± ± 1.99 1.74 2.33 2.07 1.85 2.24 98.5 98.8 98.0 99.2 97.4 96.3 ± 2.15 ± 1.98 ± 3.04 ± 1.86 ± 1.45* ± 1.94** 6262 6103 4131 294 71.3 211 735 660 76.4 14.9 2.37 2.84 31.2 42.9 107 ± 3.22* 97.5 ± 1.64* Table 2. Breeding results (* p ≤ 0.05, ** p ≤ 0.01, *** p ≤ 0.001; significance tests were done for comparison with results from 1st year) Ergebnisse der Zucht (* p ≤ 0,05, ** p ≤ 0,01, *** p ≤ 0,001; die Signifikanztests wurden im Vergleich mit dem 1. Zuchtjahr durchgeführt) Breeding year 1st 2nd 3rd Fertilization rate Hatching rate Ducklings with malformations Ducklings with motor incoordination 95% 97% 96% 64% 86%** 86%* 42% 23%** 10%*** 25% 19% 17% with malformations of the skull decreased significantly in the course of the three years from 42% in the first year to 23% in the second year and to 10% in the third year (1st year vs. 2nd year: Chi2 = 9.498, z = 3.082, p = 0.002; 1st year vs. 3rd year: Chi2 = 15.713, z = 3.964, p < 0.001). The number of (hatched) ducklings with behavioral deficits decreased insignificantly as well (1st year: 25%, 2nd year: 19%, 3rd year: 17%). Adult ducks occasionally showed skull deformations as well. These deformations are independent from incoordination of locomotion. Most frequently, perforations of the skull and encephaloceles were detected. Less frequently, additionally bone pins between 9 mm–34 mm length were found. In most cases skull perforations and bone pins were covered by the cushion of fat and connective tissue under the crest. The number of adult ducks with skull deformations decreased significantly from 62% in the first year to 36% in the second year (Chi2 = 7.176, z = 2.679, p = 0.007) (Figure 5). Discussion Based on the behavioral results we would like to stress that it is possible to classify ducks by using an easy test. Unfortunately, the distribution of the ‘righting time’ data does not allow a meaningful correlation analysis between latency to right and fat body size. Most of the data lie between 0.5 s and 5 s and thus, a necessary data range lacks. But the other results show, that the proposed ‘righting test’ is able to identify those individuals in a population that have only slightly difficulties in behavior but exhibit large fat bodies and presumably a tendency for incoordination of locomotion. These individuals bear the risk of passing a negative genetic disposition to the next generation. Now, these animals can also be selected by applying the test presented here and thus could exclude from breeding. The breeding results demonstrate the appropriateness of the test and indicate that genetic transmission may play Arch.Geflügelk. 3/2010 Mehlhorn und Rehkämper: Brain alterations and behaviour in Crested Ducks 100 Number of investigated CR Ducks without skull deformations Ducks with skull deformations 80 60 36% 40 62% 20 64% 38% 0 1st year 2nd year Figure 5. Number of CR with and without skull deformations of the 1st and 2nd breeding year Auf Schädelanomalien untersuchte Tiere aus dem 1. und 2. Zuchtjahr. a key role. This is in agreement with studies where a genetically background of these peculiarities has been assumed (REQUATE, 1959; LANCASTER, 1990). Thus, the test might help to reduce the proportion of individuals with suboptimal brains and deficits in locomotion coordination which is important for the animal welfare discussion. By analyzing the brain of CR in detail, we observed at first that an intracranial fat body seems to be a typical neuromorphological peculiarity in Crested Ducks. Presumably most of CR exhibit such a fat body. The intracranial fat body has an impact on total brain volume and on net brain volume. The increased total brain volume of CR is caused by the fat body. The size of these fat bodies rather than the existence seems to be a decisive factor for brain composition and behavior. This is in line with findings on free-living crested Ducks (“Hochbrutflugenten”) living under seminatural conditions, which have fat bodies as well and which apparently are not negatively influenced in their survival (FRAHM and REHKÄMPER, 2004). After removing of the fat body significantly smaller brains are found in CR. This is due to a reduction of several brain parts, namely the apicale hyperpallium, olfactory bulb, cerebellum and tegmentum. The olfactory bulb is strictly associated with olfaction, and the apical hyperpallium is a multimodal integration centre that serves cognitive functions and is involved in part in the visual system. These brain parts should be discussed in other contexts which are not the major topic here. The tegmentum and cerebellum are reduced as well and these findings would be in line with the observed behavioral deficits. The cerebellum can be regarded as a centre of motor coordination (ITO, 1984). It receives input from the reticular formation, the vestibular apparatus and the proprioreceptors. The tegmentum bears as well many structures that serve as motion control, for example descending motion pathways including vestibulospinal, reticulospinal and rubrospinal fibers (NIEUWENHUYS et al., 1998). Comparative neuromorphometry has elucidated a correlation between the size of a brain part and the function it serves (BENNETT and HARVEY, 1985; REHKÄMPER et al., 1988; IWANIUK and HURD, 2005) and hence suboptimal functioning of reduced structures is quite feasible. Apparently, mostly structures near the common position of the fat body are affected by reduced volume with respect to structures with a superficial position (like the olfactory bulb). Thus, the space demanding process of the fat body Arch.Geflügelk. 3/2010 207 and the space restriction from the cranium could influence the size alterations. There are many Crested Ducks with intracranial fat bodies but without other noticeable peculiarities and so this breed represents a good example for the ability of compensation and the plasticity of brains. Besides, fat bodies in CR have been discussed as being comparable to pathological lipoma found e. g. in humans (BARTELS et al., 2001). Thus, CR might serve as a model to understand genetically disposition and functional impact of such lipoma. An oversized fat body and degenerated brain structures appear to be the reason of dysfunctions and should be eliminated. Our breeding experiments have proven a relatively high degree of heritability since only few generations are needed to increase hatching rates and decrease the percentage of malformations that are found in CR. Crested Ducks have a long breeding history and their occasionally shown problems in motor coordination and the existence of intracranial fat accumulations are known since the beginning of the 20th century (KRAUTWALD, 1910). Thus, their problems should be solved, but without eliminating the whole breed. It is difficult to evaluate the consequences of the extinction of such a breed. Surely, there are a lot of fancy breeds without an obvious value for economy or nutrition of people. But, every breed is a cultural heritage and today, inbreeding and selection for more profit cause as well losing of fitness and adaptability. Species-appropriate keeping conditions and the trend to more animal welfare could put the old traditional breeds again in focus due to e.g. their adaptability or their robustness. Summary Crested Ducks (CR) are widely bred by poultry fanciers and are in focus of animal welfare activists, because they show occasionally intracranial fat bodies, incoordination of locomotion and high pre- and postnatal mortality. Here, it is shown that 24 of 26 CR bear such a fat body, but with a high variability in size (0.3% to 41% of total brain volume). A large fat body could hold responsible for motor incoordination, thus fat body size seems to be a decisive factor. A behavioural test helps to identify CR bearing a problematical fat body. Therefore, ducks were put on their back and time required to stand up was measured. The appropriateness of this test has been proven in a special breeding program. To investigate the influence of fat bodies on brain composition, an allometrically comparison of 26 CR brains with those of three uncrested duck breeds was done. Ten CR exhibited suboptimal motor coordination. CR with motor incoordination showed significantly larger fat bodies and needed significantly more time in the test than “normal” CR. Total brain volume was significantly larger in CR, but brain volume minus fat body was significantly smaller compared to reference breeds. Cerebellum, apical hyperpallium, tegmentum and olfactory bulb were significantly reduced in CR. Obviously, the behavioural deficits cannot be explained by the existence of a fat body but by functionally suboptimal cerebella and tegmenta. The relationship between fat body and reduced structures was discussed. A breeding experiment with test selected ducks resulted in increased hatching rate, decreased number of (unhatched) ducklings with skull malformations and as well decreased number of (hatched) ducks with malformations or motor incoordination. Thus, this study could be conducive to overcome the criticism of animal welfare activists without eliminating this old and traditional breed. 208 Mehlhorn und Rehkämper: Brain alterations and behaviour in Crested Ducks Crested Duck, locomotion incoordination, brain composition, intracranial fat body, righting test, breeding strategy, cerebellum paring the excellent serial sections. The support of the German Association of Poultry Breeders (BDRG) and the German foundation for the support of Young Scientists in Poultry Research (JuWiRa) is kindly acknowledged as well. Zusammenfassung References Hirnveränderungen, ihr Einfluss auf das Verhalten und Zuchtmanagement bei haubentragenden Hausenten BARTELS, T., N. KUMMERFELD, 2001: Abschlussbericht zum Forschungsauftrag 96 HS 046, Untersuchungen zur Haubenbildung bei Hausenten. Zu beziehen über die Bundesanstalt für Landwirtschaft und Ernährung, Frankfurt a.M. BARTELS, T., J. BRINKMEIER, S. PORTMANN, P. WOLF, M.E. KRAUTWALD-JUNGHANS, A. BOOS, N. KUMMERFELD, 2001: Intrakraniale Fettkörper bei Hausenten (Anas platyrhynchos f. dom). Tierärztl. Praxis. 29, 384-390. BENNETT, P.M., P.H. HARVEY, 1985: Relative brain size and ecology in birds. J. Zool. 207, 151-169. BRINKMEIER, J., 1999: Untersuchungen zur Inzidenz anatomisch-morphologischer Alterationen bei Hausenten (Anas platyrhynchos f.d.) mit Federhaube, Medical Veterinary Thesis, Tierärztliche Hochschule Hannover. CNOTKA, J., H.D. FRAHM, G. REHKÄMPER, 2006: Intrakraniale Fettkörper und ihre potentiellen Auswirkungen auf Hirnbau und Verhalten bei haubentragenden Hausenten (3 Fallstudien). Dtsch. tierärztl. Wschr. 113, 27-31. CNOTKA, J., H.D. FRAHM, A. MPOTSARIS, G. REHKÄMPER, 2007: Motor incoordination, intracranial fat bodies and breeding strategy in Crested Ducks (Anas platyrhynchos f. d.). Poult. Sci. 86, 1850-1855. CNOTKA, J., I. TIEMANN, H.D. FRAHM, G. REHKÄMPER, 2008: Unusual brain composition in Crested Ducks (Anas platyrhynchos f.d.)-including its effect on behaviour and genetic transmission. Brain Res. Bull. 76, 324-328. EBINGER, P., 1995: Domestication and plasticity of brain organization in mallards (Anas platyrhynchos). Brain Behav. Evol. 45, 286-300. FRAHM, H.D., G. REHKÄMPER, 1998: Allometric comparison of the brain and brain structures in the White Crested Polish Chicken with uncrested domestic chicken breeds, Brain Behav. Evol. 52, 292-307. FRAHM, H.D., G. REHKÄMPER, C.W. WERNER, 2001: Brain alterations in crested versus non-crested breeds of domestic ducks (Anas platyrhynchos f.d.). Poult. Sci. 80, 1249-1257. FRAHM, H.D., G. REHKÄMPER, 2004: Brain size, brain composition and intracranial fat bodies in a population of free-living crested ducks (‘Hochbrutflugenten’). Br. Poult. Sci. 45, 590-597. FRAHM, H.D., J. CNOTKA, G. REHKÄMPER, 2005: Landente ohne Haube mit neurologischen Symptomen und ihr Hirnbau. Tierärztl. Umschau. 60, 319-322. ITO, M., 1984: The Cerebellum and Neural Control, Raven, New York. IWANIUK, A.N., P.L. HURD, 2005: The evolution of cerebrotypes in birds. Brain Behav. Evol. 65, 215-230. KRAUTWALD, F., 1910: Die Haube der Hühner und Enten. Diss.med.vet., Universität Bern, Schweiz. KRUSKA, D., 1980: Changes of brain size in mammals caused by domestication. Z. Zool. Syst. Evolut.-forsch. 18, 161-195. LANCASTER, F.M., 1990: Mutations and major variants in domestic ducks. In: Crawford, R. D.: Poultry Breeding and Genetics. Elsevier, Amsterdam, 381-394. MPOTSARIS, A., G. REHKÄMPER, 2006: Automated morphometry: a new method of volume reconstruction from histological sections. J. Histotech. 29, 192. Key words Die Hausentenrasse „Landente mit und ohne Haube“ (CR) zeigt regelmäßig hohe prä- und postnatale Sterberaten, sowie Schädelanomalien, intrakraniale Fettkörper und motorische Störungen, die insgesamt als tierschutzrelevant angesehen werden („Qualzucht“). In dieser Untersuchung wird gezeigt, dass 24 von 26 Landenten einen intrakranialen Fettkörper aufweisen, dessen Größe zwischen 0,3% und 41% des Gesamthirnvolumens schwankt. Ein übergroßer Fettkörper kann für mögliche Bewegungsstörungen verantwortlich gemacht werden und als Hauptstörung angesehen werden. Es wurde ein einfacher Verhaltenstest entwickelt, der helfen soll, Tiere mit übergroßem Fettkörper leichter zu identifizieren. Dafür wurden die Tiere in Rückenlage gebracht und die benötigte Zeit bis zum Aufstehen gemessen. Der Nutzen dieses Testes wurde in einem speziellen Zuchtprogramm überprüft. Um den Einfluss des Fettkörpers auf die Hirnzusammensetzung zu untersuchen, wurden die Gehirne von 26 Landenten allometrisch mit Gehirnen von drei anderen, glattköpfigen, Hausentenrassen verglichen. Zehn der 26 CR galten als verhaltensauffällig bezüglich ihrer Bewegungskoordination. CR mit Bewegungsstörungen zeigen signifikant größere Fettkörper als ihre verhaltensunauffälligen Artgenossen und benötigen auch signifikant mehr Zeit für die Bewältigung des Verhaltenstests. Das Gesamthirnvolumen von CR ist im Vergleich zu den anderen Hausentenrassen vergrößert, nach Subtraktion des Fettkörpers jedoch signifikant kleiner. Cerebellum, Hyperpallium apicale, Bulbus olfactorius und Tegmentum sind in CR signifikant kleiner im Vergleich zu den anderen Rassen. Cerebellum und Tegmentum sind in die Motorik eingebunden und ihre Reduktion wird in Zusammenhang mit den Koordinationsproblemen diskutiert. Bei der Zucht mit positiv vorselektierten Tiere wurde eine Erhöhung der Schlupfrate erzielt. Zudem ging der Anteil an Tieren, die im Ei steckengebliebenen waren und Schädelanomalien zeigten oder nach dem Schlupf Schädelanomalien zeigten zurück. Auch reduzierte sich die Anzahl an Tieren mit Bewegungsstörungen. Mit dieser Untersuchung wird ein konstruktiver Beitrag zur Überwindung des Qualzuchtvorwurfs gegenüber den Landenten geleistet und gleichzeitig zum Erhalt der genetischen Vielfalt bei Haustieren beigetragen. Stichworte Haubenente, Bewegungsstörungen, Hirnzusammensetzung, intrakranialer Fettkörper, Umdrehtest, Zuchtmanagement, Cerebellum Acknowledgements The authors want to thank Verena Ohms for her support during the volume measures and Claudia Stolze for pre- Arch.Geflügelk. 3/2010 Mehlhorn und Rehkämper: Brain alterations and behaviour in Crested Ducks NIEUWENHUYS, R., H.J. TEN DONKELAAR, C. NICHOLSON, 1998: The central nervous system of vertebrates. Springer, Berlin, Germany. PRICE, E.O., 1999: Behavioral development in animals undergoing domestication. Appl. Anim. Behav. Sci. 65, 245-271. REHKÄMPER, G., E. HAASE, H.D. FRAHM, 1988: Allometric comparison of brain weight and brain structure volumes in different breeds of the domestic pigeon, Columba livia f. d. (Fantails, Homing Pigeons, Strassers). Brain Behav. Evol. 31, 141-149. REHKÄMPER, G., E. KART, H.D. FRAHM, C.W. WERNER, 2003: Discontinous variability of brain composition among domestic chicken breeds. Brain Behav. Evol. 61, 59-69. Arch.Geflügelk. 3/2010 209 REHKÄMPER, G., H.D. FRAHM, J. CNOTKA, 2008: Mosaic evolution and adaptive brain component alteration under domestication on the background of evolutionary theory. Brain Behav. Evol. 71, 115-126. REQUATE, H., 1959: Federhauben bei Vögeln. Eine genetische und entwicklungsphysiologische Studie zum Problem der Parallelbildungen. Z. Wiss. Zool. 162, 191-313. STEPHAN, H., G. BARON, H.D. FRAHM, 1988: Comparative size of brains and brain Components. In: Steklis, H.D., J. Erwin: Comparative Primate Biology. Alan R. Liss, New York, 1-38. Correspondence: J. Mehlhorn, C.&O. Vogt Institute of Brain Research, Heinrich-Heine-University Düsseldorf, Universitätsstrasse 1, D-40225 Düsseldorf, Germany, phone: +49 211 8112788, E-Mail address: [email protected]