3. Forage legumes and fleshy forage plants

1 Forage legumes Occurence: • Perennial or annual herbs of Fabaceae family used for their stem and leaves • Wild species on pastures • For forage: wild species and selected cultivars applied exclusively or mixed with cereals Importance: • Animal nutrition – Rich in protein and fiber – Rich in minerals: Ca and P – High content β-carotine • Pasture for honey bees • Root nodules  Rhizobium species  ability to fix athmospheic 2 N2  green manure Utilization • Grazing plants with decumbent stem (difficult to mow)  see Grasslands • Hay : plants are mow in the beginning of flowering stage 3-4 times/year) – Avoid dried leaves fall off – Drying for 2-3 days to protect ß-carotine from degradation • Dried and grinded hay comressed into pellets and cakes • Ensilage: – Silage : higher CH content which support fermentation – Haylage : drying hay to increase CH content up to 40-45% in dry weight, then ensilage (often mixed with molasses and

conserved by formic acid) 3 Antinutritive effects and compounds Bloating (distension) Freshly eaten forage legumes can be fermented easily in the intestines Water-soluble peptids of low molecular weights are released Rapid digestion by rumen microbes  slime production  frothy bloat (distension caused by foam and gases)

Effects: low O2 levels in tissues and painful spasm Taut skin, death

4 5 Saponins Amphipathic glycosides (hydrophilic and lipophilic properties) – emulsifying effect • sapo (in Latin) means soap  produce foam in the rumen • can enter into the lipid bilayer of membranes  disintegrated membranes • red blood cells are affected  haemolytic effect • low conversion rate from digestive tract • irritation of mucous membranes 6 Photosensitization (hypericosis) • Plants causing liver damage  microbially produced metabolites of chlorophyll (phyotodynamic agents)  immediate „sunburn” = dermatitis with wounds • If caused by Trifolium spp.  trifoliosis • Symptoms: inflammation of non-pigmented skin, jaundice , cholic

7 Phytoestrogenes • Isoflavanoids or coumestans • Similar to animal estrogen • Bind irreversible to estrogen receptors • Decreasing fertility – abnormal estrous cycle, abortion, sterility • Can remain in silage

(non-steroidal compounds with estrogenic activities)

8 Tannins • (Poly)phenolic molecules • Astringent taste (plant defense against herbivores)  depresses feed intake • Precipitate proteins  water insoluble complexes  decreased protein conversion rate • Fecal N excretion increases , but due to the necrosis of liver and renal tubules, total N retention can be increased as a result of decreased urinary excretion • Tannins interfere with protein and CH absorption and digestive enzymes • Animals secrete tannin-binding (mucin 9 type) proteins against tannins Toxic compounds • Cyanogenic glycosides – inhibition of cellular respiration (see seminar 4.) • Coumarin glycoside – synthesis of dicoumarol  inhibition of enzymes which activates vitamine K  abnormal blood clotting , hematomas (see seminar 7.) • Coronillin – a glucose esterified with β-nitropropionic acid molecules, symptoms are similar to Digitalis poisoning for monogastric species (vomiting, cardiac arrhythmias , seizures) (see seminar 4. and 10.)

10 Cuscuta spp. – dodder species Parasitic plants with creeping stem Quarantine pest – it’s reproduction must be controlled! Phytobesoars – plant fibers trapped in the GI tract

11 Medicago sativa – lucern, alfalfa Fabaceae – pea f. Origin: Central Asia Perennial Mainly under continental climatic conditions „al-fac-facah” (in Arabian) = father of all foods Forage for dairy cows Harvested 4 times a year

12 13 14 15 Nutritive value – antinutritive compounds Nutritive value: High protein (~20%) and fiber (20-30%) content Rich in Ca 2+ , P, vitamin K, β-carotene

Antinutritive compounds: Lucern saponins – large amount in leaves (less in new cultivars) Proteins causing frothy bloat Long term feeding can lead to: • photosensitization • bitter flavoured milk • accumulation of phytoestrogenes (coumestrol) (ruminants can digest) 16 Utilization Planted in monocultures or mixted with grasses (perennial ryegrass, orchard grass) For hay Pellet Lucerne meal Haylage

17 Other Medicago species

Medicago lupulina - black medick - in natural grasslands Medicago falcata - sickle medick - tolerates severe drought - for substitution of M. sativa - harvested one time a year, later it Medicago x varia - sand lucerne is grazed - on sandy soils, mowed one/two 18 times a year Trifolium pratense – red clover Fabaceae – pea f. Origin: Mediterranean Region Biennial or perennial (for 4-6 years) Planted for forage mainly under continental climatic condition Harvested three time a year Planted together with cereals

19 20 21 Nutritive value – antinutritive compounds

Nutritive value (compared to lucerne): Higher proportion of protein but smaller yield Produces less essential amino acids

Antinutritive compounds: Isoflavanoids – phytoestrogenes  mainly in the leaves (e.g. formononetin) Proteins causing frothy bloat Compounds inducing photosensitization

22 Trifolium incarnatum – crimson clover Fabaceae – pea f. Origin: Mediterranean Region Annual Harvested for hay (in May) before flowering stage (later dangerous for horses!) For silage Green manure With no bloating effect !

23 24 25 Nutritive value – antinutritive compounds

Nutritive value High quality hay with 25% protein content for spring (planted in autumn)

Antinutritive compounds Dry inflorescences with undigestable fibers  lumpy mass  production of phyto-trichobezoars  harvested before flowering Stem becomes more woody for summer: rapid lignification (higher fiber content, decreased digestibility)

26 Other Trifolium species

T. hybridum - alsike clover T. repens - white clover - similar nutritive value to red clover - widespread on pastures - on fresh meadows in mountains

27 Onobrychis viciifolia – sainfoin Fabaceae – pea f. Origin: Central and SE Europe Perennial Mainly on fresh meadows in mountains or planted also under more arid climatic conditions (lowlands) For substitution of lucern (similar nutritive value) More species in mountains With no bloating effect (for calves), but dried leaves fall off easly With anthelmintic effect

28 29 30 31 faeces Number of nematode eggs per one gramme gramme one per eggs nematode of Number

Comparison of faecal egg counts (arithmetic mean values) on the 4 th , 5 th and 6 th week post-infection, in the groups of lambs fed on browse plants, O. viciifolia or M. sativa . Results of statistical analysis based on analysis of variance on repeated measurements * P < 0.05; ** P < 0.01. 32 Other forage legumes

Melilotus albus – white sweet clover Low protein content, stem early becomes woody with age (rapid lignification) Coumarin glycosides  abnormal blood clotting (see seminar 7.) With a slight bloating effect when fresh Green manure

33 Lotus corniculatus – Can be found in pastures – birdsfoot trefoil applied for hay

Securigera (Coronilla)

varia – crown vetch 34 Deseases caused by forage legumes: a summary Diseases and symptoms Ages, causes Plant species Frothy bloat: distension, water-soluble peptids of low lucerne, red clover, white digestive flatulence molecular weights, saponins clover, alsike clover, white sweet clover Trifoliosis / hypericosis liver damages caused directly lucerne, red clover, white secondary photosensitiza- by poisonous compounds clover, alsike clover, white tion: dermatitis with wounds, from legumes; reaction of sweet clover jaundice, colic, paralysis photodynamic agents to (UV) light  dermatitis

Decreased fertility, abnormal phytoestrogens lucerne, red clover estrous cycle, abortion, (isoflavonoids, coumestans): sterility with effects similar to that of animal estrogen

Hemolysis, frothy bloat saponins: disintegrate cell lucerne, red clover, white membranes clover, alsike clover, white sweet clover Constipation, diarrhea, tannins: interference with sainfoin, kidney vetch necrosis of liver and renal protein and CH absorption tubules and digestive enzymes; but 35 anthelmintic effect Deseases caused by forage legumes: a summary

Diseases and symptoms Ages, causes Plant species HCN toxicity: histotoxic cyanogenic glycosides: white clover, birdsfoot trefoil hypoxia, blood (and skin) of inhibitionof cellular respiration cherry colour; clotting of blood is slow; pink mucous membranes Abnormal blood clotting, coumarin glycosides: white sweet clover hematoma inhibition of vitamine K production Cardiac arrhythmias, coronillin: stimulation of crown vetch vomiting, seizures, inhibitory ganglia of heart death Loosing weight despite the phytobezoar: indigestible crimson clover, kidney vetch, fact of sufficient feeding, plant materials (mainly dodder species nausea, vomiting, gastric fibres); too many weeds on outlet obstruction, the pasture, or in the hay perforation, abdominal pain

36 Fleshy forage plants • Large biomass • High water content (75-90%) – large proportion of water is

unavailable for metabolic optional surface processes tightly loosely available • Low proportion of proteins, fatty adsorbed adsorbed for acids and fibers metabo- • Easily digestable compounds: starch and lism sugar (5-25%) • Can be eaten freshly (after shredding) • With laxative effect • Digestive problems may occure when frozen, decayed, moldy or soil-contamined forages are applied 37 I. MODIFIED TAPROOT: • Beta vulgaris subspecies crassa varietas altissima – Amaranthaceae – sugar beet • Beta vulgaris ssp. crassa var. crassa – Amaranthaceae – fodder beet • Daucus carota – Apiaceae – carrot II. MODIFIED STEM: • Solanum tuberosum – Solanaceae - potato • Helianthus tuberosus – Asteraceae – Jerusalem artichoke III. FLESHY LEAVES: • Brassica oleracea – Brassicaceae – forage kale IV. FLESHY FRUITS: • Cucurbita pepo – Cucurbitaceae - marrow 38 Species with modified taproot • Nutritive value: 80-90% water, 10-20% dry weight (mainly CH) • Increase milk production for dairy cows • Fresh or fermented taproot (after shredding) • During storage: must protect from freezing and infection of Clostridium spp. (bacteria causing diarrhea) • Covered by soil particles – must wash! - • Species with high nitrate ion (NO3 ) level!

39 Beta vulgaris The ancestor of current beet types: Beta vulgaris spp. maritima – sea beat

Beet cultivars planted currently for forage, or sugar production were raised between the XIV. and XVII. centuries in Western Europe.

40 Beta vulgaris ssp. vulgaris var. altissima – sugar beet Amaranthaceae – amaranth f. Planted mainly in Europe and E-Asia Biennial Mainly for sugar production – many by-products for forage Whole plants are rarely used as fresh forage for cattle

41 Beetroot = taproot + hypocotyl (ebryonic stem) 42 43 44 Nutritive value – antinutritive compounds Nutritive value: • Rich in saccharose (12-20%) • Contains many lysine and vitamine A

Antinutritve compounds: • Consumption of too much sugar (beet)  lactic acidemia • Too much sugar in rumen  decreased pH  shifts in species abundances in rumen microbial community  digestive problems • Accumulation of nitrate and oxalate ions (for poisoning, see  seminar 7.) • Betain – can be secreted in the milk (beet/fish 45 taste) 46 Beet tops with leaves

• The first by-product of sugar production process • Must be washed • For ruminants • Freshly or can be ensilaged

47 48 49 By-products of sugar production

• Beet pulp (after sugar has been extracted) – for cows, (pigs, horses): freshly or ensilaged • Filter cake (dried pulp) – high energy and fiber content (for monogastric species): soaking in water or grinded when dry • Pellets – dried, compressed pulp • Molassess (40-50% sugar) – mixing with forages, triggers ensilage or applied with carbamid (urea)

50 1,25-1,35t of sugar

5-7t of tops with leaves

0,5t of molasses 4t of beet pulp

10t of sugar beet

51 Other Beta species Beta vulgaris ssp. vulgaris var. crassa – fodder beet

Planted in small farmlands for forage only Mainly for dairy cows A fresh forage for winter Nutritive value and antinutritive compounds similar to sugar beet Higher yield

52 53 54 Other Beta species

Beta vulgaris ssp. vulgaris var. cicla – chard

Beta vulgaris ssp. vulgaris var. conditiva – beetroot 55 Other species with modified taproot Daucus carota ssp. sativus – carrot (Apiaceae )

• Mainly for human consumption, but for horses, young, ill and breeding animals (vitamine supply !) • High sugar content - palatable • Many ß-carotene and K +, Ca 2+ , Mg 2+ , Se, vitamine C • Used to enrich carotene content in milk and butter • Dried for rodents or fermented • Several antinutritives but in low amounts (high level in wild taxa)

• Accumulation of nitrate ions! 56 57 58 Fleshy forages with a modified stem

Tuber  underground stem enlagred for storing nutrients (mainly starch) Axillary buds on its surface

59 Solanum tuberosum – potato Solanaceae – nightshade f.

Origin: S America (Andes mountains) Perennial Thousands of cultivars Forage for cattle and pig For human consumption (starch, alcohol, chips…) GMO potato

60 61 62 Nutritive value – antinutritive compounds

Nutritive value: • Starch content: 80% (in the dry matter) • Low protein content: 10% (in the dry matter) • Contains vitamine A, B, C, K For pigs cooked  small feed conversion rate of raw starch Raw tubers for cattle

Antinutritive compounds: • Protesae (trypsin) inhibitors chaconine • Glycoalkaloids: solanin and chaconine (aglycon: solanidin): thermostable but water-soluble compounds

• In green tubers: 0,02%, in fruits: 1%, in germs: 1,7% 63 Solanine poisoning Inhibition of acetylcholinesterase  CNS symptoms Disrupts cell membranes Vomiting, salivation, mucosal inflammation, diarrhea Dizziness, paralysis of muscles, cardiac arrhythmia, death Higher solanine level: tubers exposed to sunlight , mechanical damage Solanine level: 2-15mg/100g (kept in darkness), 80-100mg/100g (green tubers) Application of moldy, damaged and sprouting tubers must be avoided! 64 65 Humans are more sensitive to the toxic effects of chaconine and solanine than other mammals.

66 67 Other species with modified stem Helianthus tuberosus – Jerusalem artichoke (Asteraceae )

Compared to potato, with a lower proportion of CH (15-16%) High levels of inulin (a polyfructan): 75-80% in dry weight, no starch Pigs and ruminants (game) prefer (sweet taste), good feed conversion Undigestable for humans – sweetener for diabetics Antinutritive compounds: solanine- like compounds in germinating and young plant

Stem can be ensilaged 68 69 Forage plants with fleshy leaves

Fleshy leaves, high biomass The highest proportion protein content in leaves Many vitamine B, C, and carotine Low fiber proportion, high water content

70 Brassica oleracea convar. acephala – forage kale Brassicaceae – mustard f. Origin: SW Europe Biennial One of the latest green forages in vegetation period Proteins : 15-20% Fresh even in January (vitamin C supply ) Mainly for cattle Can be ensilaged For decoration 71 72 Antinutritive compounds • Glucosinolates (see oilseed rape! seminar 2.) • S-methyl-L-cysteine sulfoxide (SMCO): digestion dysfunctions, loss of apetite, lesions in liver and kidneys, anemia-linked symptoms (lower milk yield and growth rate) • Accumulation of nitrate ions (for poisoning, see seminar 7.) • Frothy bloat  limited quantities should be applied

73 VFI – voluntary feed intake LWG – liveweight gain

74 75 Brassica oleraceae ssp. sabauda – Savoy cabbage

Seasonally grazed Can regrow after it nibbled With similar nutritive and antinutritive

characteristics than forage kale 76 Forage plants with fleshy fruits Cucurbita pepo – marrow Cucurbitaceae – marrow f. Origin: N America Very high water content: 90-95% Many cultivars with a high level of carotenes  improve milk production Seed contains oils (30%) – oilseed cake for forage Cucurbitacins – bitter phytosterols causing digestive problems : salivation, vomiting, diarrhea Difficult to store (fungal infections) 77 78 79 Cucurbitacin poisonings

80 cytotoxic effects  targets of human cancer researches

sweet cultivars with no substantial cucurbitacin contents, but backcrossing can often be observed with wild pepo species

81