The browser wars ( )
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1 The browser wars ( ) Marcus Clauss Clinic for Zoo Animals, Exotic Pets and Wildlife, Vetsuisse Faculty, University of Zurich, Switzerland Wildlife Digestive Physiology Course Vienna 2013
2 The term "browser wars" is the name given to the competition for dominance in the web browser marketplace - the struggle between Internet Explorer and Netscape Navigator during the late 1990s, and the growing threat which Mozilla Firefox poses to Internet Explorer from 2004 onward. (Wikipedia)
3 But why is the web-browser called browser?
4 browser = animal that eats (mainly) browse? especially as opposed to grazer = animal that eats (mainly) grass?
5 Did you know that there are grazing and browsing mites? Siepel H, de Ruiter-Dijkman EM (1993) Feeding guilds of oribatid mites based on their carbohydrase activities. Soil Biol Biochem 25:
6 Did you know that there are grazing and browsing carnivorous fish? Lechanteur YARG, Griffiths CL (2003) Diets of common suprabenthic reef fish in False Bay, South Africa. Afr Zool 38:
7 Coupled with Hofmann s term concentrate selector, the word browser has become a synonym for an organism selectively feeding on highly digestible material.
8
9 until 1970: All ruminants are similar and function as cattle and sheep do.
10 Digestive adaptations
11 Digestive adaptations
12 Digestive adaptations
13 Digestive adaptations
14 What comparative digestive physiology can offer to domestic ruminant research Understanding where ruminants came from in evolutionary terms from
15 What comparative digestive physiology can offer to domestic ruminant research Understanding where domestic ruminants came from among the ruminants from Agnarsson et al. (2008)
16 What comparative digestive physiology can offer to domestic ruminant research Understanding where domestic ruminants came from among the ruminants and where they might be taken to in the future from Agnarsson et al. (2008)
17 (from Nickel-Schummer-Seiferle 1967)
18 (from Nickel-Schummer-Seiferle 1967) Rumen
19 (from Nickel-Schummer-Seiferle 1967) Rumen
20 (from Hofmann & Schnorr1982)
21 Reticulum Rumen (from Nickel-Schummer-Seiferle 1967)
22 Reticulum Rumen (from Nickel-Schummer-Seiferle 1967)
23 (from Nickel-Schummer-Seiferle 1967)
24 (from Nickel-Schummer-Seiferle 1967) Omasum
25 (from Nickel-Schummer-Seiferle 1967) Omasum
26 Omasum Abomasum (from Nickel-Schummer-Seiferle 1967)
27 (from Nickel-Schummer-Seiferle 1967)
28
29 Stratification of rumen contents: cattle-type gas layer fibre mat : large particles of low density longish, fibrelike particles liquid layer grass small particles of high density from Clauss et al. (2003)
30 Stratification of rumen contents: cattle-type gas layer fibre mat : large particles of low density liquid layer small particles of high density from Hummel et al. (2009)
31 Stratification of rumen contents: cattle-type from Clauss et al. (2010)
32 Rumen of addax - a grazer from Clauss et al. (2009)
33 Stratification of rumen contents Rumen dors. Rumen ventr. Reticulum Dry matter (%) Cattle Moose from Clauss et al. (2010)
34 Testing stratification by ultrasound - cattle from Tschuor & Clauss (2008)
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36 Stratification and rumen papillation dorsal Atrium ventral from Clauss, Hofmann et al. (2009)
37 Digestion and Sorting Rumen: Fermentation (from Grau 1955)
38 Digestion and Sorting Reticulum: Sorting Rumen: Fermentation (from Grau 1955)
39 Digestion and Sorting Reticulum: Sorting Rumen: Fermentation (from Grau 1955)
40 Functional density of particles
41 Functional density of particles un-fermented ingesta particle: entangles in fibre mat
42 Functional density of particles un-fermented ingesta particle: entangles in fibre mat size reduction by rumination/ attachment of bacteria
43 Functional density of particles un-fermented ingesta particle: entangles in fibre mat size reduction by rumination/ attachment of bacteria fermentation activity = gas production => adhesion of gas bubbles => updrift/low density
44 Functional density of particles un-fermented ingesta particle: entangles in fibre mat size reduction by rumination/ attachment of bacteria fermentation activity = gas production => adhesion of gas bubbles => updrift/low density fermented ingesta particles: high density
45 Sorting in the reticulum un-fermented particles, low density, large fermented particles, high density, small (from von Engelhardt & Breves 2000)
46 Ruminants always rest in sternal recumbency
47 (there may be pigs in space, but no sheep on the moon!)
48 Sorting by density... fermentation = gas production gas adhesion = updrift fermented particle no gas bubbles = high density
49 Sorting by denisty... flotation and sedimentation only works in a fluid medium
50 the fluid must be removed... high moisture content in the reticulum (from Nickel-Schummer-Seiferle 1967)
51 the fluid must be removed... high moisture content in the reticulum it would be difficult for the abomasum to work against the dilution (from Nickel-Schummer-Seiferle 1967)
52 the fluid must be removed... high moisture content in the reticulum it would be difficult for the abomasum to work against the dilution therefore the omasum removes fluid (from Nickel-Schummer-Seiferle 1967)
53 roughage ration concentrate ration Distinct rumen contents stratification Less distinct rumen contents stratification (from Lauwers 1973)
54 roughage ration concentrate ration Distinct rumen contents stratification Less distinct rumen contents stratification Distinct dry matter gradient between rumen contents and reticulum outflow Small dry matter gradient between rumen contents and reticulum outflow (from Lauwers 1973)
55 roughage ration concentrate ration Distinct rumen contents stratification Less distinct rumen contents stratification Distinct dry matter gradient between rumen contents and reticulum outflow Small dry matter gradient between rumen contents and reticulum outflow Omasum with high resorption capacity Omasum with low resorption capacity (from Lauwers 1973)
56 until 1970: All ruminants are similar and function as cattle and sheep do.
57
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62 B R O W S E R G R A Z E R
63 Ruminant feeding types (Hofmann) from Hofmann (1989)
64 Ruminant feeding types (Hofmann) from Hofmann (1989)
65 Ruminant feeding types (Hofmann) from Hofmann (1991)
66 Ruminant feeding types (Hofmann) from Hofmann (unpubl.)
67 Ruminant feeding types (Hofmann) from Hofmann (1985)
68 > 460 citations (and counting)
69 Ruminant feeding types (Hofmann) dicot (Hofmann 1989) monocots browser grazer
70 Ruminant feeding types (Hofmann) dicot (Hofmann 1989) monocots browser grazer selective? un-selective?
71 Feeding types dicot monocot
72 Feeding types selective un-selective dicot monocot
73 Feeding types selective un-selective dicot monocot
74 Feeding types from Van Soest (1996)
75 Ruminant feeding types (Hofmann) from Hofmann (1989)
76 Ruminant feeding types (Hofmann) from Hofmann (1989)
77 Ruminant feeding types (Hofmann) from Hofmann (1989)
78 Ruminant feeding types (Hofmann) from Hofmann (1989)
79 Don t lose perspective!
80 Don t lose perspective! is slower than...
81 Don t lose perspective! a slow car?
82 Don t lose perspective!
83 Don t lose perspective! contains less alcohol than...
84 Don t lose perspective! an alcohol-free beverage?
85 Don t lose perspective!
86 Don t lose perspective! eats a diet of lower fibre content than...
87 Don t lose perspective! a concentrateselector?
88 Crude fibre in rumen contents (Drescher-Kaden & Seifelnasr 1977) (%dry matter)
89 Crude fibre in rumen contents (Drescher-Kaden & Seifelnasr 1977) (%dry matter) 20 % Area 1 20 %
90 Crude fibre in rumen contents (Drescher-Kaden & Seifelnasr 1977) (%dry matter) 20 % Area 1 20 % 24 % Area 2 34 %
91 Roe deer select against fibre content (Lignocellulose) (Duncan et al. 1998) available forage 24% Lignozellulose (%dry matter)
92 Roe deer select against fibre content (Lignocellulose) (Duncan et al. 1998) available forage 24% Lignozellulose selected diet 18% Lignocellulose (%dry matter)
93 Do diets of grazers and browsers really differ? Species Crude fiber (% dry matter) Giraffe (Giraffa camelopardalis) Okapi (Okapia johnstoni) Moose (Alces alces) White-tailed deer (Odocoileus virginianus) Buffalo (Syncerus caffer) Waterbuck (Kobus ellipsiprymnus) NDF (% dry matter) from Clauss & Dierenfeld (2008)
94 Do diets of grazers and browsers really differ? Crude fibre in rumen contents (%DM) from Hofmann (1989) and Woodall (1992)
95 Do diets of grazers and browsers really differ? Crude fibre in rumen contents (%DM) 23% 16% 20% 18% 31% 25% 21% 21% 21% 31% 24% from Hofmann (1989) and Woodall (1992)
96 Differences between grass and browse Sugar Starch Pectin Hemicellulose Cellulose [% DM] [% DM] [% DM] [% DM] [% DM] Grass Browse from Robbins (1993)
97 Differences between grass and browse Sugar Starch Pectin Hemicellulose Cellulose [% DM] [% DM] [% DM] [% DM] [% DM] Grass Browse ? from Hofmann (1989)
98 Differences between grass and browse 100% 80% NDF 60% 40% HC C ADL? 20% 0% Grass Browse Twigs Herbs Legumes from Hofmann (1989)
99 Ruminant feeding types (Hofmann) from Hofmann (1985)
100 Ruminant feeding types (Hofmann)... based on anatomical and dietary observations... from Hofmann (1985)
101 Ruminant feeding types (Hofmann) from Hofmann (1989)
102 Ruminant feeding types (Hofmann) from Hofmann (1989)
103 Ruminant feeding types (Hofmann) anatomical parameters from Hofmann (1989)
104 Ruminant feeding types (Hofmann) behavioral parameters from Hofmann (1989)
105 Feeding bout frequency from Hummel et al. (2006)
106 Differences between grazers and browsers: anatomy... and physiology? from Kay et al. (1984)
107 Differences between grazers and browsers: anatomy... and physiology? ok from Kay et al. (1984)
108 Differences between grazers and browsers: anatomy... and physiology? ok ok from Kay et al. (1984)
109 Differences between grazers and browsers: anatomy... and physiology? ok ok ok from Kay et al. (1984)
110 Differences between grazers and browsers: anatomy... and physiology? ok ok ok ok from Kay et al. (1984)
111 Differences between grazers and browsers: anatomy... and physiology? ok ok ok ok ok from Kay et al. (1984)
112 Differences between grazers and browsers: anatomy... and physiology? ok ok ok ok ok ok from Kay et al. (1984)
113 Differences between grazers and browsers: anatomy... and physiology? ok ok ok ok ok ok from Kay et al. (1984)
114 Differences between grazers and browsers: anatomy... and physiology? ok ok ok ok ok ok from Kay et al. (1984)
115 Pyhsiological postulates based on anatomical observations without experimental data!
116 from Hofmann (1989)
117 Additionally, Hofmann claimed that the most influential factor for all physiological processes body size was less important than the feeding type classification!
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125 Ad libitum intake of grass hay in zoo ruminants Grazer Intermediate Giraffe OM intake (kg) Body weight (kg) from Foose (1982)
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127 Moose study: energy offered as concentrates 70 energy offered/animal (MJ ME/d) Facility from Clauss (2000)
128 Moose study: energy offered as concentrates 70 energy offered/animal (MJ ME/d) calculated requirement Facility from Clauss (2000)
129
130 Problems with hay acceptance Giraffe Fox (1938), Gradwell (1976), Kloeppel (1976), Altmann (1978), Gorgas et al. (1978), Brancker (1980), Foose (1982), Franz et al. (1984), Gutzwiller (1984), Hofmann and Matern (1988), Matern and Kloeppel (1995) Moose Baines (1965), Landowski (1969), Heptner and Nasimowitsch (1974), Bo and Hjeljord (1991), Schwartz (1992), Schwartz and Hundertmark (1993), Shochat et al. (1997) Mule deer Cahart (1943), Doman and Rasmussen (1944), Nagy et al. (1969), Schoonveld et al. (1974) Roe deer Dissen (1983) Chinese water deer Hofmann et al. (1988) Duiker Cowan (1982), Luginbuhl et al. (1991), Van Soest et al. (1995) Reindeer Eriksson and Schmekel (1962), Kurkela (1976), Valtonen et al. (1983) Eland Hofmann (1973, p. 40), Miller et al. (2010) Kudu Miller et al. (2010)
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144 No difference in ingesta retention time between the feeding types. All differences in ingesta retention can be explained by body weight. according to Gordon and Illius (1992, 1994)
145 Mean retention time (particles) = 15.3 * BM for all feeding types From Illius & Gordon (1992); no particle size given; data e.g. from Foose (1982)
146 Ruminant feeding types (Hofmann) from Hofmann (1989)
147 Ruminant feeding types (Hofmann) from Hofmann (1989)
148 Illius & Gordon s (1992) equation reliably predicts retention time in grazers Species BM MRT (h) (kg) calculated measured Mouflon Mountain sheep Cattle data from Udén et al. (1982), Baker & Hobbs (1987), Behrend et al. (2004)
149 Illius & Gordon s (1992) equation overestimates retention time in browsers Species BW MRT (h) (kg) calculated measured Roe deer Okapi Giraffe data from Clauss et al. (1998), Clauss and Lechner-Doll (2001), Behrend et al. (2004); particle size < 2 mm
150
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