Metabolic Rate and Kleiber's Law

Kleiber's law, named after Max Kleiber's biological work in the early 1930s, is the
observation that, for the vast majority of animals, an animal's metabolic rate scales to the
power of the animal's mass. Symbolically: if q0 is the animal's metabolic rate, and M the
animal's mass, then Kleiber's law states that q0 ~ M. Thus a cat, having a mass 100 times that
of a mouse, will have a metabolism roughly 32 times greater than that of a mouse. In plants,
the exponent is close to 1.
The first accurate measurements of body mass versus metabolic rate in 1932 shows
that the metabolic rate R for all organisms follows exactly the 3/4 power-law of the body
mass, i.e., R

M3/4. This is known as the Kleiber's Law. It holds good from the smallest

bacterium to the largest animal. The relation remains valid even down to the individual
components of a single cell such as the mitochondrion, and the respiratory complexes (a
subunit of the mitochondrion) as shown in Figure 02. It works for plants as well.
Hukum Kleiber, bernama setelah bekerja biologis Max Kleiber di awal 1930-an,
adalah pengamatan bahwa, untuk sebagian besar hewan, timbangan tingkat metabolisme
hewan dengan kekuatan massa hewan. Secara simbolis: jika q0 adalah tingkat metabolisme
hewan, dan M massa hewan, maka hukum Kleiber menyatakan bahwa q0 ~ M. Jadi kucing,
memiliki massa 100 kali dari mouse, akan memiliki metabolisme sekitar 32 kali lebih besar
dari mouse. Pada tumbuhan, eksponen dekat 1.
Pengukuran yang akurat pertama massa tubuh terhadap tingkat metabolisme pada
tahun 1932 menunjukkan bahwa tingkat R metabolisme untuk semua organisme berikut
persis 3/4 kekuatan-hukum massa tubuh, yaitu, R M3 / 4. Hal ini dikenal sebagai Hukum
Kleiber itu. Ini berlaku baik dari bakteri terkecil sampai hewan terbesar. Hubungan tetap
berlaku bahkan ke masing-masing komponen sel tunggal seperti mitokondria, dan kompleks
pernafasan (subunit dari mitokondria) seperti yang ditunjukkan pada Gambar 02. Ia bekerja
untuk tanaman juga.

Concept of metabolic efficiency (ME)

This limit to blood flow considerations is problematic when claims are made that the
theoretical models also are relevant to things without blood flow, like bacteria and coral.

Moreover, despite assurances that the equation models metabolic efficiency, the term
metabolic efficiency (ME) appears nowhere in the equation as expounded upon by its major
proponents. When the term is added to the exponent (which becomes (4ME-1)/4ME, where
ME is metabolic efficiency), and the exponent is or , ME is 100 or 89%. Efficiencies like
these are not found in nature unless thermogenesis is included as part of metabolic rate. This
removes the WBE version of Kleiber's Law, which the metabolic theory of ecology rests on,
from any biological relevance whatsoever. The efficiency that is purported to be modeled is
actually assumed.
Attempts to understand the metabolic rate of a multi-cellular organism (field
metabolic rate, that includes the activity of the organism) are couched in terms of the product
between average basal metabolic rate, and number of cells. This, plus capillary terminal size
invariance, leaves the equation open to the criticism that it cannot possibly account for spikes
in metabolic rate needed for motor activity. Too much blood would be required. This
intimates that as proposed and popularly handled, the equation does not have the relevance to
biology claimed, and is based upon assumptions that are not part of the equation, like
fractality.
In plants, according to a paper in 2006 in Nature, the exponent of mass is close to 1.
Mathematically this is not possible since the implication is that ME is greater than 100% in
the case of plants. The key problem is the nature of metabolic energy and the extent of what
is considered metabolism. The problem is most clearly noticeable in the unit term for
metabolic rate, i.e., calories/s. Calories are a measure of heat energy. This leads to the idea
that thermogenesis is part of metabolism, Kleiber's original treatment, and rules out that
metabolism is all about chemical energy, not heat energy. The picture is further obfuscated
when the idea of respiratory metabolism is introduced to refine and limit the definition of
metabolism such that oxygen consumption and synthesis of ATP are its ultimate factors. The
data from study of oxygen consumption metabolic rates in cells in vitro suggests that the
exponent is not only far less than , but also becomes negative for things less than one gram
in size. Furthermore, glycogenesis is excluded from metabolic consideration on this model
since glycogenesis is not included in the respiratory chain, and is itself a reduction reaction
not strictly dependent upon the proximity of certain molecules and atoms delivered by
capillaries and vibrating from Brownian motion. Energy is required for glycogenesis, and the

blood does not deliver energy, just the ingredients for endergonic reactions. The energy
comes from redox coupling, what ME is all about.
ME amends these problems, and describes the metabolic rate in watts. Metabolic rate
becomes the rate at which a biomass recharges so that its degeneration is prevented, and its
organization is perpetuated. ME is a ratio of the rate of reduction reactions necessary for the
maintenance, growth, replication and behavior of the biomass, to the rate of availability of
energy captured and expended by that biomass. ME is a statement of redox coupling
efficiency. ME excludes thermogenesis as part of metabolism, consequently. A graph of
Kleiber that includes ME, with ME as the X axis, metabolic rate as the Y axis, and a different
curve for each mass, reveals a picture of the relation of biomass to metabolic rate that
suggests all of evolution took place at less than 45% ME. Furthermore, this graph models the
relation between basal metabolic rate and field metabolic rate, where the latter is the rate for
the organism while the former is the rate for its cells. The graph clearly depicts that, contrary
to WBE and Savage, biomass for an organism has no effect on basal metabolic rate (BMR);
that what influences BMR is the ME of the organism, not its mass. The organism determines
ME, and that ME is the same for it and for its cells.
The failure of WBE to parse Kleiber to understand the nature of aging is due to their
mishandling of its mathematics. This mishandling results from failure to limit considerations
of metabolism to strictly electrochemistry, and to exclude heat generation. Both of these
things follow from failure to consider the role of variable ME on that biomass. If metabolic
rate (MR) is taken as the recharge rate in watts of an electrolytic biomass, then MR is directly
related to the longevity of that biomass. The equation, when graphed, suggests that for
creatures operating at over 25% ME, the organism lives longer than its cells. What this
indicates is that the organism has a source of new cells that are not initially a part of it,
undifferentiated stem cells for example, especially in creatures over one gram mass. Aging
appears to be the result of antagonism between BMR and FMR (field metabolic rate), given
fluctuations in ME. These fluctuations are driven predominantly by alterations in the
denominator of the ratio ME, where that denominator represents the availability of food
sources.
Batas ini pertimbangan aliran darah yang bermasalah ketika klaim yang dibuat bahwa
model teoritis juga relevan dengan hal-hal tanpa aliran darah, seperti bakteri dan karang.
Selain itu, meski ada jaminan bahwa efisiensi metabolisme model persamaan, efisiensi

metabolik jangka (ME) tidak ada dalam persamaan yang diuraikan oleh pendukung utama.
Ketika istilah ini ditambahkan ke eksponen (yang menjadi (4me-1) / 4me, di mana ME adalah
efisiensi metabolisme), dan eksponen adalah atau , ME adalah 100 atau 89%. Efisiensi
seperti ini tidak ditemukan di alam kecuali thermogenesis dimasukkan sebagai bagian dari
tingkat metabolisme. Hal ini menghilangkan versi WBE dari Kleiber Hukum, yang teori
metabolis ekologi bersandar pada, dari relevansi biologi apapun. Efisiensi yang konon
dimodelkan sebenarnya diasumsikan.
Upaya untuk memahami tingkat metabolisme organisme multisel (tingkat
metabolisme bidang, yang meliputi aktivitas organisme) yang ditulis dalam istilah produk
antara rata-rata tingkat metabolisme basal, dan jumlah sel. Ini, ditambah terminal kapiler
ukuran invarian, daun persamaan terbuka untuk kritik yang tidak mungkin menjelaskan
lonjakan tingkat metabolisme yang dibutuhkan untuk aktivitas motorik. Terlalu banyak darah
akan diperlukan. Hal ini mengisyaratkan bahwa seperti yang diusulkan dan populer ditangani,
persamaan tidak memiliki relevansi biologi mengklaim, dan didasarkan pada asumsi-asumsi
yang bukan bagian dari persamaan, seperti fraktalitas.
Pada tumbuhan, menurut sebuah makalah pada tahun 2006 di Nature, eksponen massa
dekat dengan 1. Secara matematis hal ini tidak mungkin karena implikasinya adalah bahwa
ME lebih besar dari 100% dalam kasus tanaman. Masalah utama adalah sifat energi
metabolisme dan tingkat apa yang dianggap metabolisme. Masalahnya adalah yang paling
jelas terlihat dalam jangka satuan untuk tingkat metabolisme, yaitu, kalori / s. Kalori adalah
ukuran energi panas. Hal ini menyebabkan ide bahwa thermogenesis adalah bagian dari
metabolisme, pengobatan asli Kleiber itu, dan aturan bahwa metabolisme adalah semua
tentang energi kimia, tidak panas energi. Gambar selanjutnya dikaburkan ketika gagasan
metabolisme pernapasan diperkenalkan untuk memperbaiki dan membatasi definisi
metabolisme sehingga konsumsi oksigen dan sintesis ATP adalah faktor utamanya. Data dari
studi tingkat metabolisme konsumsi oksigen dalam sel in vitro menunjukkan bahwa eksponen
tidak hanya jauh kurang dari , tetapi juga menjadi negatif untuk hal-hal yang kurang dari
satu gram dalam ukuran. Selanjutnya, glikogenesis dikeluarkan dari pertimbangan
metabolisme pada model ini karena glikogenesis tidak termasuk dalam rantai pernapasan, dan
itu sendiri merupakan reaksi reduksi tidak sepenuhnya tergantung pada kedekatan molekul
tertentu dan atom disampaikan oleh kapiler dan bergetar dari gerak Brown. Energi yang
dibutuhkan untuk glikogenesis, dan darah tidak memberikan energi, hanya bahan-bahan

untuk reaksi endergonik. Energi yang berasal dari kopling redoks, apa ME adalah semua
tentang.
ME merubah masalah ini, dan menggambarkan tingkat metabolisme dalam watt.
Tingkat metabolisme menjadi tingkat di mana biomassa mengisi sehingga degenerasi yang
dicegah, dan organisasi diabadikan. ME adalah rasio laju reaksi reduksi diperlukan untuk
pemeliharaan, pertumbuhan, replikasi dan perilaku biomassa, dengan tingkat ketersediaan
energi ditangkap dan dikeluarkan oleh biomassa itu. ME adalah pernyataan efisiensi redoks
kopling. ME tidak termasuk thermogenesis sebagai bagian dari metabolisme, akibatnya.
Grafik Kleiber yang meliputi ME, dengan ME sebagai sumbu X, tingkat metabolisme sebagai
sumbu Y, dan kurva yang berbeda untuk setiap massa, mengungkapkan gambaran hubungan
biomassa untuk tingkat metabolisme yang menunjukkan semua evolusi berlangsung kurang
dari 45% ME. Selain itu, grafik ini model hubungan antara tingkat metabolisme basal dan
tingkat metabolisme lapangan, di mana yang terakhir adalah tingkat untuk organisme
sementara mantan adalah tingkat untuk sel-sel. Grafik jelas menggambarkan bahwa,
bertentangan dengan WBE dan Savage, biomassa untuk organisme tidak berpengaruh pada
tingkat metabolisme basal (BMR); bahwa apa yang mempengaruhi BMR adalah ME
organisme, bukan massanya. Organisme menentukan ME, dan bahwa AKU adalah sama
untuk itu dan untuk sel.
Kegagalan WBE untuk mengurai Kleiber memahami sifat penuaan adalah karena
kesalahan penanganan mereka matematika nya. Ini kesalahan penanganan hasil dari
kegagalan untuk membatasi pertimbangan metabolisme ketat elektrokimia, dan untuk
mengecualikan panas. Kedua hal ini mengikuti dari kegagalan untuk mempertimbangkan
peran ME variabel pada biomassa itu. Jika tingkat metabolisme (MR) diambil sebagai tingkat
resapan dalam watt dari biomassa elektrolit, maka MR secara langsung berkaitan dengan
umur panjang biomassa yang. Persamaan, ketika digambarkan, menunjukkan bahwa untuk
makhluk beroperasi pada lebih dari 25% ME, organisme hidup lebih lama dari sel-sel. Ini
menunjukkan bahwa organisme memiliki sumber sel-sel baru yang awalnya tidak menjadi
bagian dari itu, sel-sel induk berdiferensiasi misalnya, terutama dalam makhluk lebih dari
massa gram. Penuaan tampaknya merupakan hasil dari pertentangan antara BMR dan FMR
(tingkat metabolisme lapangan), mengingat fluktuasi ME. Fluktuasi ini didorong terutama
oleh perubahan dalam penyebut dari rasio ME, di mana penyebut yang mewakili ketersediaan
sumber makanan.

Optimal foraging theory

Optimal foraging theory (OFT) is a model that helps predict how an animal behaves when
it's searching for food. Although obtaining food provides the animal with energy, searching for
and capturing the food require both energy and time. The animal wants to gain the most benefit
(energy) for the lowest cost during foraging, so that it can maximize its fitness. OFT helps predict
the best strategy that an animal can use to achieve this goal.
OFT is an ecological application of the optimality model. This theory assumes that the
most economically advantageous foraging pattern will be selected for in a species
through natural selection.[1] When using OFT to model foraging behavior, organisms are said to
be maximizing a variable known as the currency, such as the most food per unit time. In addition,
theconstraints of the environment are other variables that must be considered. Constraints are
defined as factors that can limit the forager's ability to maximize the currency. The optimal
decision rule, or the organisms best foraging strategy, is defined as the decision that maximizes
the currency under the constraints of the environment. Identifying the optimal decision rule is the
primary goal of the OFT.

The Three Types of Functional Responses

There are three general types of curves expected in various predator-prey situations
that show by Holling. That is :
Type I is a linear relationship, where the predator is able to keep up with increasing
density of prey by eating them in direct proportion to their abundance in the environment. If
they eat 10% of the prey at low density, they continue to eat 10% of them at high densities.
The dotted line indicates a maximum consumption rate that some authors attach to Type I
foraging.
Type II describes a situation in which the number of prey consumed per predator
initially rises quickly as the density of prey increases but then levels off with further increase
in prey density.
Type III resembles Type II in having an upper limit to prey consumption, but differs in
that the response of predators to prey is depressed at low prey density.

Two factors dictate that the functional response should reach a plateau. First, the predators
may become satiated (i.e., their stomach completely filled), at which point their rate of
feeding is limited by the rate at which they can digest and assimilate food. Second, as the
predator captures more prey, the time spent handling and eating the prey lowers the searching
time. Eventually, the predator reaches the minimum time it takes to search, capture and
consume. The predator cannot find prey and eat it any faster. The consumption rate cannot
increase when the ability of the predator to catch and eat is at a maximum. C.S. Holling
described this relationship between search time, handling time, and consumption rate by a
simple expression known as the disc equation (Spring, 2001).

Hollings disk equation

Optimal diet models are largely derivatives of Hollings disk equation which gives a
technique for calculating the rate of return from foraging behavior and takes into account the
real-life variables of rate of reward collection, search time and handling time. What I like best
about these models is that you can actually measure the components with real animals and
test the theoretical predictions.
Holling actually had blindfolded students collect sandpaper disks that he arrayed on a
table. He timed their movements and analyzed their strategies to generate this equation that
describes the behavior of a forager. Because Holling used an empirical approach he did not
actually derive the equation; rather, he fit an equation to his data. derive the disk equation by
starting with a simpler formulation:

RATE = Energy gained in foraing/(time searching + time handling) put into symbols, this
equation reads:
R = Ef/(Ts + Th)

R = (le - s)/(1 + lh)

where: s = cost of search per unit time l = rate of encounter with items h = handling time e =
energy gained per encounter In theory, a forager finds the behavior that maximizes its return
under this model. Basically, the forager needs to assess/remember the four elements of this
equation (Stephens and Krebs, 1986) .

REFERENCES
Stephens, D. W. and J. R. Krebs. 1986. Foraging theory. USA : Princeton University Press.
Spring. 2001. Journal Holling Disc Equation: Student Manual Biology 1240.
Begon, M., J. L. Harper & R. Colin, 1996. Ecology: individuals, populations and
communities, Blackwell Scientific Publications, London.
Krebs, JR. & NB. Davies, 1993. Behavioral ecology: an evolutionary approach. 3rd ed.
Blackwell Scientific Publications, London
Krebs JR. 1989. Ecological methodology. New York. Harper Collins Publisher.

Odum, PE., 1993. Dasar-dasar ekologi, diterjemahkan oleh Samingan,T. & B. Srigandono,
Gadjah Mada University Press.