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27 Role of Minerals in Halophyte Feeding to Ruminants SALAH A. ATTIA-ISMAIL Desert Research Center, Matareya, 11753 Cairo, Egypt 1 INTRODUCTION Halophyte plants are those that grow naturally in saline environment. Halophytes grow in many arid and semi-arid regions around the world and are distributed from coastal areas to mountains and deserts. Saline conditions may include either salt- affected soil or saline water irrigation. Economically speaking, the most productive halophytic plant species yield from 10 to 20 ton/ha of biomass when irrigated with seawater [Glenn et al., 1999]. Also, the productivity of cultivated halophytes, as compared with traditional crops, is high [Jaradat, 2003]. The average area of salt-affected soils might be calculated on the basis of FAO averages to be 450 million hectares. That means we are talking in about 4–5 billion tons of biomass. Halophytic plants have many uses: They can be used as animal feeds, vegetables, and drugs, as well as for sand dune fixation, wind shelter, soil cover, cultivation of swampy saline lands, laundry detergents, paper production, and so on [El Shaer and Attia-Ismail, 2002]. Feed shortage especially in coastal areas has led to the exploration and exploitation of marginal resources such as halophytic plants. Halophytes can, then, play an important role in the welfare of people in such regions. As an animal feed component, halophytes are promising because they have the potentiality of being good feed resource. The potentialities of halophytes as animal feed components were recognized as early as the 1880s [Hutchings, 1965]. Yet, this potentiality does not go far because several constraints are limiting and have to be worked out. One of these constraints is the high contents of salts and conse- quently the high mineral composition. Trace Elements as Contaminants and Nutrients: Consequences in Ecosystems and Human Health, Edited by M. N. V. Prasad Copyright # 2008 John Wiley & Sons, Inc. 701
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Role of Minerals in Halophyte Feeding to Ruminants

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Page 1: Role of Minerals in Halophyte Feeding to Ruminants

27 Role of Minerals in HalophyteFeeding to Ruminants

SALAH A. ATTIA-ISMAIL

Desert Research Center, Matareya, 11753 Cairo, Egypt

1 INTRODUCTION

Halophyte plants are those that grow naturally in saline environment. Halophytesgrow in many arid and semi-arid regions around the world and are distributed fromcoastal areas to mountains and deserts. Saline conditions may include either salt-affected soil or saline water irrigation.

Economically speaking, the most productive halophytic plant species yield from10 to 20 ton/ha of biomass when irrigated with seawater [Glenn et al., 1999].Also, the productivity of cultivated halophytes, as compared with traditional crops,is high [Jaradat, 2003]. The average area of salt-affected soils might be calculatedon the basis of FAO averages to be 450 million hectares. That means we aretalking in about 4–5 billion tons of biomass. Halophytic plants have many uses:They can be used as animal feeds, vegetables, and drugs, as well as for sanddune fixation, wind shelter, soil cover, cultivation of swampy saline lands, laundrydetergents, paper production, and so on [El Shaer and Attia-Ismail, 2002].

Feed shortage especially in coastal areas has led to the exploration and exploitationof marginal resources such as halophytic plants. Halophytes can, then, play animportant role in the welfare of people in such regions.

As an animal feed component, halophytes are promising because they have thepotentiality of being good feed resource. The potentialities of halophytes as animalfeed components were recognized as early as the 1880s [Hutchings, 1965]. Yet,this potentiality does not go far because several constraints are limiting and haveto be worked out. One of these constraints is the high contents of salts and conse-quently the high mineral composition.

Trace Elements as Contaminants and Nutrients: Consequences in Ecosystems andHuman Health, Edited by M. N. V. PrasadCopyright # 2008 John Wiley & Sons, Inc.

701

Page 2: Role of Minerals in Halophyte Feeding to Ruminants

2 ASH AND MINERAL CONTENTS OF HALOPHYTES

The high content of ash is a typical characteristic of halophytes. Mineralcontents differ in halophytes from those of regular forages. This has led to scientificconcerns of with regard to what extent this may affect their feeding value toanimals. Also, the mineral compositions of the high ash contents of halophyteshave been controversial: Do they support animals’ requirements of certain minerals,do they might exceed the requirements to the extent that they may represent a poison-ing threat to animals? Mineral contents of halophytes vary considerably. Thevariations in mineral contents of halophytes depend on several factors. Amongthe many factors that cause the variations are the plant species, stage of growth,and season.

Table 1 shows the ash and mineral contents of different parts of some halophytesaround the world. Halophytes may represent a good source of minerals to ruminantanimals. High ash contents of halophytes, unless associated with nondigestible com-ponents such as silica, may compensate for mineral deficiency usually seen inanimals in areas depending mainly on rangeland grazing as well as desert andcoastal areas. Table 1 shows that ash contents of halophytes vary for whole plants,maturity stage, and plant part. It runs up to more than 40% of plant materials.Plant parts differ also in their ash contents. Leaves and twigs of Atriplex repanda,for instance, have 23.1%, whereas the fruits contain 32.65% (Table 1); this represents41% increment in ash content. Salicornia bigelovii spikes have 66% more ash thanthe stem. The stem of Sueda foliosa has 117% more ash than leaves.

Stage of plant maturity affects ash contents as well. Pappophorum caespitosumis an example. Its ash contents decreased after fructification by 98.5%. Trichloriscrinita is a contrast case where ash contents increased after fruitification by 152%.The variability in ash contents might be due to the physiological distribution andpools of minerals in different plant parts. Physiological and biochemical processesthat take place in plants vary according to the plant species, and this mayexplain the contrast in the percentage of ash content increment or decrement due tothe process of fruitification between both of Pappophorum caespitosum andTrichloris crinita.

3 FACTORS AFFECTING MINERAL CONTENTSOF HALOPHYTES

Halophyte plants either grow naturally in saline environment (salt-affected soils orunder the circumstances of the irrigation with saline water) or need be cultivated.Either way has its impacts on ash contents and mineral composition of the plants.Minerals in halophytes differ widely according to the particularities of the environ-ment they grow within. Type of soil (saline or sodic), level of salinity (in eithersoil or irrigation water), mineral profile of the soil, plant species, season, stage ofgrowth, part of the plant, and so on, are among factors affecting ash and mineralcontents of halophytes. Mineral contents are, therefore, affected accordingly.

702 ROLE OF MINERALS IN HALOPHYTE FEEDING TO RUMINANTS

Page 3: Role of Minerals in Halophyte Feeding to Ruminants

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Page 4: Role of Minerals in Halophyte Feeding to Ruminants

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Page 5: Role of Minerals in Halophyte Feeding to Ruminants

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Page 6: Role of Minerals in Halophyte Feeding to Ruminants

4 SALT-AFFECTED SOILS

Saline soils are the soils that have high percentages of salts. Saline soils have an elec-trical conductivity at the saturation of soil extract of more than 4 dS/m [FAO, 1988].Salt-affected soils are distributed all over the world. Salt-affected soils are classifiedinto two main classes according to the type of salts present in soils: saline soils(solonchak) and sodic soils (solonetz) [Szabolcs, 1974]. FAO [2007] estimated thatthe solonchak soils area varies from 260 to 340 million hectares and are mostlyfound in Saharan Africa, East Africa, Central Asia, Australia, and South America.The solonetz soils are associated with solonchak soils and are estimated to bearound 135 million hectares (http://www.fao.org/ag/agl/agll/wrb/wrbmaps/htm/).Saline soils were classified into three main groups according to the dominantcations: the sodium-dominated soils, the magnesium-dominated soils, and thecalcium-dominated soils (http://www.fao.org/DOCREP/003/Y1899E/y1899e00.htm#toc). In each class of these soils the ratios of cations differ.

The most common salts present in saline soils are sodium, potassium, calcium andmagnesium in the form of chlorides, sulfates or bicarbonates. The sodium and chlo-ride are present in the highest percentage particularly in highly saline soils. The mostcommon sodic salts present in sodic soils are the carbonates [FAO, 1988] particularlysodium.

5 IRRIGATION WITH SALINE WATER

Saline water contains large amounts of dissolved salts. For water to be saline, itshould include more than 1000 ppm dissolved salts; this value can be as high as35,000 ppm for typical seawater. Saline water usually goes along with saline soilsor in some cases independently of saline soils. Saline water varies in its salinity con-tents according to the source (seawater, underground water, or drainage water). Salinedrainage waters may contain toxic levels of trace elements as well as the macro-minerals [Retana et al., 1993]. The minerals in saline water differ according to itssource. Brackish water may contain toxic levels of certain minerals according to theeffluent. Major elements of seawater are Ca, 419 ppm; Mg, 1304 ppm; Na, 10,710;K, 390; Cl, 19,350 (http://www.palomar.edu/oceanography/salty_ocean.htm).

6 SALINITY LEVEL

Salinity level either in irrigation water or contained in soils is a major determinantfactor in plant growth and, therefore, in ash contents and mineral profile in plants.Brown et al. [2006] found that salinity level affects nutrient uptake in Spartinaalterniflora. Daoud et al. [2001] mentioned that includer halophytes may accumulatesalt and, therefore, increase mineral concentrations at increasing salinity level, whileexcluder halophytes may also accumulate salt but at lower rates. Table 2 shows thateffect clearly. However, it appears that different parts of the plant respond differently.

706 ROLE OF MINERALS IN HALOPHYTE FEEDING TO RUMINANTS

Page 7: Role of Minerals in Halophyte Feeding to Ruminants

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Page 8: Role of Minerals in Halophyte Feeding to Ruminants

At higher salinity levels that reached seawater salinity, sodium concentrationsincreased in shoots while potassium concentrations decreased in roots (Table 2).Noticeably, the salinity level did not affect the concentrations of magnesium orcalcium in their experiments. The distribution of ash and, therefore, mineral contentsin different plant pools might be a mechanism by which the plants adapt to a salineenvironment [Gorham, 1996].

Parida et al. [2004] used Bruguiera parviflora to test the effect of salinity leveltolerance, by adding increasing levels of sodium chloride, on some mineral contentsof different parts of this mangrove (Table 3). The increase in sodium chloride concen-trations led to increases in the concentrations of both of Na and Cl ions in all testedparts of the plant. Khan [2001] reported that the increase in sodium chloride uptakecounteracts the uptake of potassium, calcium, and manganese in some plants.

Certain mineral ratios are affected by salinity level as a consequence of increasingsalinity level like the K/Na, Ca/Na, and many other ratios.

7 PLANT SPECIES

Halophyte plant species differ in their ability to accumulate minerals [Reboreda andCacador, 2007]. The ability of halophytes to accumulate minerals depends on themechanism by which halophytes distribute minerals in different plant compartments.Minerals in halophytes are mainly accumulated in the roots with small quantitiestranslocated to the stems and leaves [Windham et al., 2003]. Halophytes havehigher potentiality to phytoextract minerals from the soil than do glycophytes[Jordan et al., 2002].

Legumes, generally, have different mineral profile (being high in calcium,potassium, magnesium) than grasses which tend to be higher in manganese andmolybdenum when grown under the same conditions.

8 MINERAL ROLE IN RUMINANT NUTRITION

Minerals, in general, either major or minor, play a crucial role in the lives of animalsand affect their production to a greater extent. Minerals have four vital functions inthe bodies of animals. They have a structural function in bones and other structuraltissues of the body, physiological function in body fluids as electrolytes, regulatoryfunction as they regulate several metabolic processes in the body, and catalytic func-tion when they enhance the enzyme activities [Underwood and Suttle, 1999].

9 RECOMMENDED MINERAL ALLOWANCES

Minerals required by animals for proper functioning of the body and for the optimalproduction are either macro- or micro-minerals. Macro-minerals are known to be Ca,

708 ROLE OF MINERALS IN HALOPHYTE FEEDING TO RUMINANTS

Page 9: Role of Minerals in Halophyte Feeding to Ruminants

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965

.41+

0.7

137.

1+

1.8

4.35

+0.

717

.05+

0.9

7.28

+0.

110

015

.81+

0.7

7.86

+0.

72.

91+

0.6

61.5

4+

0.3

187.

0+

0.8

5.69

+0.

321

.07+

0.4

21.9

1+

0.3

200

16.7

0+

0.1

3.81

+0.

32.

26+

0.7

88.3

4+

0.5

190.

5+

0.5

2.39

+0.

514

.01+

0.7

26.7

0+

0.6

400

21.6

7+

0.2

2.71

+0.

23.

63+

0.5

298.

41+

0.3

220.

4+

1.5

1.48

+0.

912

.63+

0.5

38.5

4+

0.8

Roo

t0

7.21

+0.

525

.45+

0.4

2.74

+0.

620

5.68

+0.

863

2.1+

1.8

65.9

0+

0.9

28.4

0+

0.5

6.85

+0.

710

014

.47+

0.3

17.7

8+

0.1

2.89

+0.

315

9.55

+0.

497

0.9+

1.1

51.1

0+

0.5

37.0

0+

0.5

18.3

7+

0.3

200

17.2

5+

0.5

15.0

3+

0.9

2.45

+0.

814

2.87

+0.

538

3.9+

0.9

28.1

0+

1.2

33.0

7+

0.3

24.9

5+

0.5

400

26.9

6+

0.8

11.2

0+

0.5

2.38

+0.

311

6.78

+0.

627

7.8+

0.6

14.5

2+

0.7

11.9

6+

0.7

31.4

7+

0.3

709

Page 10: Role of Minerals in Halophyte Feeding to Ruminants

P, Na, Cl, Mg, K, and S, while the micro-minerals are Fe, I, Cu, Zn, Mo, Mn,Co, and Se.

Recent studies have revealed some of the requirements of goats and sheep in dryareas. NRC [1981] recommended mineral requirements for goats and for sheep[NRC, 1985]. Table 4 shows some recommended mineral requirements for goatsand sheep. Clear variations are present when matching mineral supply of halophytesto the requirements of these animal species. Depending on halophytes as the onlymineral source for the animals may result in either mineral deficiency of someelements, toxicity of some others, or malabsorption of some other elementsbecause of the presence of different proportions of them (e.g., Ca/P, K/Na,Ca/Na ratios). However, mineral requirements differ according to several conditionssuch as the season (summer versus winter, especially for the desert animal) and thephysiological state (pregnant, lactating, growing, etc.).

10 MINERALS DEFICIENCY IN HALOPHYTE INCLUDED DIETS

Ash contents of halophytes may limit animal feed intake and digestibility coefficientsof ruminant diets. Therefore, some conservation is placed on the use of these plantspecies in livestock ratios. Adequate trace mineral intake and absorption is requiredfor a variety of metabolic functions including immunity, reproduction, and so on.

Animals, especially those inhabiting desert areas, suffer from high environmentalheat stress. A typical characteristic of animals in desert areas is to dissipate heatthrough several adaptive mechanisms. One of them is to sweat. Sweating in such

TABLE 4. Recommended Mineral Requirements of Sheep and Goats

NRC

Mineral Lactating Goats, 1981 Sheep, 1985

Na (% of DM) 0.09–0.18 0.09–0.18Cl (% of DM) — —Ca (% of DM) 0.20–0.80 0.20–0.82P (% of DM) 0.20–0.40 0.16–0.38Mg (% of DM) 0.12–0.18 0.12–0.18K (% of DM) 0.50–0.80 0.50–0.80S (% of DM) 0.14–0.26 0.14–0.26I (ppm) 0.10–0.80a 0.10–0.80Fe (ppm) 30–50a 30–50Cu (ppm) 10–20a 7–11Mo (ppm) 0.50–1.0a 0.5Co (ppm) 0.10–0.20a 0.1–0.2Mn (ppm) 20–40a 20–40Zn (ppm) 20–33a 20–33Se (ppm) 0.10–0.20a 0.1–0.2

aRick Machen, Texas Agriculture Extension Service.

710 ROLE OF MINERALS IN HALOPHYTE FEEDING TO RUMINANTS

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animals [Farid, 1989] is accompanied by the excretion of salts from the body. Thisauthor cited (from MacFarlane et al. [1963]) that camels’ sweat is rich in bicarbonates(pH 8.2–8.5) and is particularly high in potassium, four times as much as sodium.When animals are exposed to high environmental temperatures, the sweating rateincreases as a result (about 0.5 kg/p day in sheep and goats). Thus, higher concen-trations of urea, sodium, potassium, and chloride might be found in the sweat ofanimals [Farid, 1989]. Sweat was found to contain different amounts of Mg, Na,K, Ca, and Cl in humans [Verde et al., 1982].

Animals in these areas suffer from mineral deficiency or are exposed to high con-centrations of various minerals in either feedstuffs or drinking water (if brackish wateris used) or both. For example, Walker [1980] found that the concentration of calciumand potassium is usually higher than that of other minerals, with the average being1–1.5% in Southern African browsers. The deficiency results from either excessivesweating or through the feeding on halophyte plants that are either deficient incertain minerals or that may have imbalanced mineral ratios. The opposite appliesfor mineral toxicity. Figure 1 shows the symptoms of mineral deficiency inanimals fed on Atriplex halimus. The clear calcium deficiency is manifested inthese pictures [Khattab, 2007]. Figure 2 also shows lambs in a state of convulsion,exhibiting retracted head on chest with extension of limbs, while Fig. 3 showslambs in a state of muscular deformities abduction at the foreleg. Figure 4 alsoshows lambs in a state of paralysis of the hind legs; these lambs exhibit incoordina-tion of movement, tremors, convulsion, reluctance to move, aimless movement, andwandering.

Goats browsing naturally growing halophytes in Mexico [Haenlein and Ramirez,2007] were deficient in minerals intake. They found a significant low supply of Mg,Cu, Mn, and Zn from the range plants browsed by goats compared to the requirementsof these minerals. On the other hand, sheep selected different range plants and haddeficiency in Ca, Mg, K, Cu, and Mn. Similarly, phosphorus deficiencies inacacias were reported by Vercoe [1986] and Craig et al. [1991] which led to an imbal-ance in the Ca/P ratio in foliage. Panicum turgidum was fed to camels in the south-eastern region of Egypt [Badawy, 2005]. Mineral composition of this halophyte is

Figure 1. Manifestation of calcium deficiency in lambs fed Atriplex halimus. Courtesy ofDr. Khattab, Desert Research Center, Cairo, Egypt.

10 MINERALS DEFICIENCY IN HALOPHYTE INCLUDED DIETS 711

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presented in Table 5. The major problem encountered was the extreme low Ca/P ratiowhich reached 1:19 in wet season and 1:32 in dry season. This ratio is far below thatgenerally recommended (2:1) for domestic animals. They found a similar trend in Mgconcentrations. It tended to be higher in range during dry season than in wet season.

Figure 2. Lambs in a state of convulsion, retracted head on chest with extension of limbs.Courtesy of Dr. Khattab, Desert Research Center, Cairo, Egypt.

Figure 3. Lambs in a state of a muscular detormities adduction at the foreleg and abduction atthe foreleg. Courtesy of Dr. Khattab, Desert Research Center, Cairo, Egypt.

Figure 4. Lambs in a state of paralysis of the hind legs. These lambs exhibit incoordination ofmovement, tremors, convulsion, reluctance to move, aimless movement, and wandering.Courtesy of Dr. Khattab, Desert Research Center, Cairo, Egypt.

712 ROLE OF MINERALS IN HALOPHYTE FEEDING TO RUMINANTS

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The levels of Mg and K in Panicum turgidum were inadequate compared with thedietary requirements of ruminants recommended by NRC [1981].

11 EXCESSIVE MINERALS IN LIVESTOCK RATIONSIN DRY AREAS

Some halophytes attain higher levels of some trace or major elements than do othertraditional feedstuffs [Attia-Ismail, 2005]. It seems that some of their mineral concen-trations are within normal ranges to the extent that they may support normal animalphysiological functions. On the other hand, excessive concentrations may reach toxiclevels to the animals. Certain mineral ratios are skewed. For instance, the calcium–phosphorus ratio present in most halophytes does not match that required byanimals to support normal functions. In general, excessive sodium, chloride, andpotassium contents of halophytes may have several drawbacks to the animal health.The capacity of the animal body to store excess electrolytes or to excrete in the feces isvery little [Masters et al., 2007]. Marai et al. [1995] found that animals have low rectaltemperature and high water retention when exposed to higher levels of sodium.

12 EFFECT OF HALOPHYTES FEEDING ON MINERALUTILIZATION

Excess or deficient mineral intake may adversely affect diet and mineral utilization.However, the degree of absorption (bioavailability) of minerals may affect the stateof mineral utilization by animals. The bioavailability of minerals is an importantfactor that has to be taken into consideration. It varies, however, among different min-erals because of different conditions. Haenlein and Ramirez [2007] found that thedegree of absorption of minerals differ and that it was low for Cu, Mn, Mg, Zn,Ca, and Fe.

Attia-Ismail et al. [2003] fed sheep a mixture of salt plants and found that Caintake (mg/kg BW0.75) of salt plant-fed animals was higher (Table 6) (P , 0.05)than that of a control group. In a previous study [Fahmy et al., 2001], animals con-sumed more salt plants (SP) than control (hay). Therefore, the amount of Ca intake

TABLE 5. Mineral Content in Panicum turgidum as BasalDiet for Camels

Mineral Wet Season Dry Season DS þM

Ca% 1.69 1.62 1.60P% 0.09 0.05 0.04Mg% 0.23 0.27 0.26Na% 0.35 0.42 0.40K% 0.23 0.31 0.30

Source: Badawy [2005].

12 EFFECT OF HALOPHYTES FEEDING ON MINERAL UTILIZATION 713

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of the SP group and, hence, the balance were higher (P , 0.05) than control. Yet,serum Ca concentrations (mg%) of the SP group was almost similar to that of thecontrol group. Sodium (Na) intake of salt plant-fed groups was, logically, higherthan control group (Table 6).

Fahmy [1998] found Na intake to vary from 805 to 1507 mg/kg BW0.75 forsilages made of a mixture of salt plants; a range close to that was obtained in thework carried out by Attia-Ismail et al. [2003]. The Na balance in the SP-fedanimals was, however, lower than that of the control group. The excreted Na inthis experiment might be higher for the SP-fed animals than for the control group.Serum Na did not differ (Table 6). Potassium utilization followed the same trendas Na did except that serum K of control was higher than the SP-fed animals.

13 EFFECT OF MINERALS ON RUMEN FUNCTION

The role of mineral elements in rumen microbial activities has not generally receivedenough attention in work dealing with mineral metabolism in ruminants. Minerals,VFA, lactate, and glucose are the primary solutes in ruminal fluid. Short-chainfatty acids contribute to the osmotic pressure in the rumen. Major minerals contribute

TABLE 6. Mineral Utilization of Sheep Fed a Mixture of HalophyticPlants [Attia-Ismail et al., 2003]

Halophyte-Fed Sheep Control Group

Calcium Utilization

Ca intake (mg/BW0.75) 914a 484b

Ca balance (mg/BW0.75) 914a 341b

Serum Ca (mg%) 8.2a 10.9a

Sodium Utilization

Na intake (mg/BW0.75) 1546a 1068b

Na balance (mg/BW0.75) 252a 316a

Serum Na (mg%) 421a 431a

Potassium Utilization

K intake (mg/BW0.75) 1138 1049K balance (mg/BW0.75) 8b 61a

Serum K (mg%) 25.5a 61b

Zinc Utilization

Zn intake (mg/BW0.75) 45.2a 31.2b

Zn balance (mg/BW0.75) 40.4a 28.6b

Serum Zn (mg%) 1.22a 0.85b

a,bValues bearing different superscripts in the same row differ significantly( p , 0.05).

714 ROLE OF MINERALS IN HALOPHYTE FEEDING TO RUMINANTS

Page 15: Role of Minerals in Halophyte Feeding to Ruminants

more than short-chain fatty acids to rumen osmolalrity [Bennink et al., 1978].Usually, the osmotic pressure in the rumen is rather constant (250–280 mOsmol/kg),but it can increase up to 350–450 mOsmol/kg after feeding certain diets (e.g.,lucerne pellets [Bennink et al., 1978]). It was concluded that the increment in potass-ium concentrations caused a rise in rumen osmolalrity to about 400 mOsmol/kg[Warner and Stacy, 1965]. Barrio et al. [1991] observed a closer relationshipbetween osmolarity of rumen fluid and potassium concentration than betweenosmolalrity and sodium concentration in sheep. However, the increase in rumenosmotic pressure is followed by an increase in blood osmotic pressure [Dobsonet al., 1976]. Camels fed on rations containing Atriplex nummularia showed higherblood Na values (448.3 and 454.3 mg/100 ml) than those of other tested groups[Kewan, 2003]. Khattab [2007] had the same results when Atriplex halimus was fedto sheep (Table 7).

Rumen volume increased when osmotic pressure was increased through sodiumchloride increasing doses [Lopez et al., 1994]. Yet, it was concluded that theosmotic pressure effect is mainly exerted on the rate and extent at which waterenters or leaves the rumen pool rather than on the absolute size of the rumen poolitself. It is, therefore, that the washout of the digesta from the rumen increases.Harrison et al. [1975] suggested that such an increase in the flow of digesta to theduodenum would increase the efficiency of microbial synthesis in the rumen,thereby increasing the flow of protein and starch to the duodenum.

Remond et al. [1993] found that the increase in osmolality after NaCl injectionslightly decreased NH3 absorption. The increase in osmolality from 260 to 420mOsmol/liter was accompanied by a drop in pH [Hogan, 1961].

14 EFFECT OF MINERALS ON FEED INTAKE

Some researchers suggest that the Na and K chlorides play a controlling effect on feedintake. Bergen [1972] studied the possible role of osmotic pressure in altering

TABLE 7. Blood Minerals Content of Barki Ewes Fed the Experimental Diets DuringPregnancy and Lactation Periods (mean+++++SE)

Mixture of Atriplex halimus and Acaciasaligna

Items Control Fresh Silage

Na (mmol/liter) 148.68+3.5c 173.15+5.03a 167.85+2.38b

K (mmol/liter) 4.68+0.79b 5.30+0.43a 167.85+2.38b

Ca (mg/dl) 11.57+0.31a 9.70+0.57b 9.01+0.39b

Zn (mg/dl) 96.33+0.04a 78.65+0.05b 77.03+0.05b

Mg (mg/dl) 2.53+0.36a 1.53+0.02b 1.26+0.04b

Cu (mg/dl) 117.5+6.20a 53.3+3.05b 47.5+5.05b

Se (mg/dl) 11.58+4.01b 22.60+6.50a 19.87+5.93a

a– cValues bearing different superscripts in the same column differ significantly (P , 0.05).

14 EFFECT OF MINERALS ON FEED INTAKE 715

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voluntary feed intake when rumen osmolalrity was artificially elevated to morethan 400 mOsm/kg with NaCl or NaAc just before feeding. He suggested that theshort duration of elevated osmotic pressure when feeding roughages is not animportant factor in controlling feed intake. Kato et al. [1979] reached the same con-clusion when they infused sheep with potassium chloride or sodium chloride duringmeals. On the other hand, when Ternouth and Beattie [1971] infused NaCl, KCl, orsodium salts of organic acids into fistulated sheep, they found that the reduction infood intake is related to osmolalrity of the added fluid and that the reduction was moresignificant when potassium chloride was infused than with sodium chloride. Theyexplained that on the basis of the probable relation of the slower absorption of potassiumcompared with sodium. An elevation in osmotic pressure in the rumen is sensed bythe wall of the reticulo-rumen to inhibit feed intake [Carter and Grovum, 1990].Forbes [1995] concluded that the relative importance of osmolalrity as a satiety factorin ruminants is still uncertain. However, the results of the effect of increased saltintake are inconsistent.

Fahmy et al. [2001] fed a mixture of sun-dried salt plants [22.5% as Zygophyllumalbum, 22.5% as Halocnemum strobilaceum, 45% as Tamarix mannifera, and 10%molasses (w/w)] to growing sheep. The mixture represented 40% of the totalration. The salt-plant-fed group had a higher feed intake than that of the Berseemhay-fed group. Total dry matter intake was comparable and did not differ significantlyamong animal groups. Khattab [2007] fed a mixture of fresh and silage of bothAtriplex halimus and Acacia saligna to sheep lambs. Feed intakes were 1.57 forcontrol, 1.45 for fresh atriplex and acacia, and 1.62 kg/hr/day for ensiled atriplexand acacia. Sodium intakes were 10.81, 24.13, and 21.54 while potassium intakeswere 22.62, 15.74, and 11.69 g/hr/day, respectively.

15 EFFECT OF MINERALS ON WATER INTAKEAND NUTRIENT UTILIZATION

The salt content in halophytes can influence the animal’s water requirement becauseadditional water is required to excrete their high salt content. The high salt levels ofthe foliage increase the water requirements of grazing animals [Wilson, 1974].Gihad [1993] reported that drinking water increased by 61.4% when sheep werefed Atriplex halimus instead of clover hay. Puri and Garg [2001] observed that sup-plementation of sodium chloride increased voluntary water intake, rumen volume,and outflow rate from the rumen. Rapid influx of water to neutralize osmoticpressure swells the ruminal papillae and can pull patches of the ruminal epitheliuminto the rumen by stripping the internal surface layers of the rumen wall fromthe underlying layers, as illustrated vividly in histological studies by Eadie andMann [1970]. Swingle et al. [1996] observed that lambs fed halophyte diets con-sumed up to 110% more water per day and 50% more water per kilogram of drymatter intake.

716 ROLE OF MINERALS IN HALOPHYTE FEEDING TO RUMINANTS

Page 17: Role of Minerals in Halophyte Feeding to Ruminants

16 EFFECT OF MINERALS ON MICROBIAL COMMUNITYIN THE RUMEN

It is proposed that the increased salt intake and, hence, the osmotic pressure of therumen may affect the microorganism population and metabolism. Increasingosmotic pressure in the rumen is believed to be unfavorable to the growth of protozoa.Also, the increased rates of outflow are believed to contribute to decreased protozoalpopulation [Warner and Stacy, 1977]. The artificial rise of rumen osmotic pressure[Bergen, 1972] up to 400 mOsmol/kg caused cellulose digestion to be inhibited.Shawket et al. [2001] reported that increased salt intake increased dilution rate ofthe rumen fluid phase and negatively affected the protozoal population. Kewan[2003] reported that the total protozoal count (�103) per milliliter was significantlyhigher (P � 0.05) for camels fed on rations that contained Atriplex nummularia thanfor those fed on Acacia saligna and treated rice straw rations. Total rumen protozoalcount in camels fed on berseem hay ration were significantly (P � 0.05) higher thanthose fed on the other experimental rations.

In conclusion, halophytic plants may be used as a feed component in ruminant rationsin spite of the precautions placed on their use in animal feeds. However, if these plants aretreated the right wayaccording to the limitations of each plant species, they would providea plausible feed resource for ruminants in areas like arid, semi-arid, and coastal areas.This way the feed shortage in these areas will be relieved.

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720 ROLE OF MINERALS IN HALOPHYTE FEEDING TO RUMINANTS