Plants have various adaptations to maintain thermal balance with their environment. They absorb radiation and lose excess heat through evapotranspiration and convection. Temperature affects photosynthesis, with different plant species having optimal temperatures. It also influences plant growth processes like flowering through the calculation of degree days. Plants withstand extreme temperatures through adaptations like changes to cell membranes and solute concentrations that allow supercooling or survival of high temperatures through morphological and metabolic adaptations.
Plants have various adaptations to maintain thermal balance with their environment. They absorb radiation and lose excess heat through evapotranspiration and convection. Temperature affects photosynthesis, with different plant species having optimal temperatures. It also influences plant growth processes like flowering through the calculation of degree days. Plants withstand extreme temperatures through adaptations like changes to cell membranes and solute concentrations that allow supercooling or survival of high temperatures through morphological and metabolic adaptations.
Plants have various adaptations to maintain thermal balance with their environment. They absorb radiation and lose excess heat through evapotranspiration and convection. Temperature affects photosynthesis, with different plant species having optimal temperatures. It also influences plant growth processes like flowering through the calculation of degree days. Plants withstand extreme temperatures through adaptations like changes to cell membranes and solute concentrations that allow supercooling or survival of high temperatures through morphological and metabolic adaptations.
Chapter 7 PLANT ADAPTATIONS TO THE ENVIRONMENT II:
THERMAL, MOISTURE, AND NUTRIENT ENVIRONMENTS.
PLANTS AND THE THERMAL ENVIRONMENT Plants are constantly absorbing short-wave and long-wave radiation from the surrounding environment. THERMAL ENERGY BALANCE Plants maintain a thermal balance through evaporation and convection. Leaf size, shape and the stomatal opening/closing control influence these processes. he energy absorbed by plants per unit of time is referred to as the plant!s net radiation balance, "n. Plants absorb and reflect solar radiation, and absorb short-wave radiation and emit long-wave radiation. he difference between the two is the net radiation balance. "n # $ % & % 'C % ()* Less than +, of the absorbed radiation is used in photosynthesis and stored in chemical bonds, M. )nergy is also used in heating the plant tissues and raising the temperature of the boundary layer, S. Convection 'C* and evaporation 'E* dissipate energy into the environment- evaporation includes transpiration and direct evaporation. # latent heat of vaporization- energy re.uired to transform one unit of li.uid water to vapor. ranspiration and evaporation occur in plants/ evapotra!p"rat"o. )vaporation occurs from the surface of leaves. 0eat is dissipated by evaporation and transpiration. 1ir temperature directly affects evaporation and transpiration. Tra!p"rat"o is the result of stomatal conductance and vapor pressure deficit. Cove#t"o depends on the temperature difference between the plant and the surrounding air. 2actors influencing heat loss by convection/ 3ifference between leaf and air temperatures. Conductance of the boundary layer. 4ind removes the warm air of the boundary layer and increases the difference between the leaf and the atmosphere- the boundary layer conductance increases with the movement of air. he size and shape of the leaf affects the conductance of the boundary layer. he ratio of surface to volume affects heat convection. &mall, lobed leaves are more effective at heat e5change than are larger, less lobed leaves. Plants must replace the water lost in evapotranspiration. 1ir temperature and wind velocity impose limits on the ability of plants to dissipate e5cess heat energy. 4ater lost must be replaced- therefore, precipitation patterns have a direct effect on the plant!s energy balance. THERMAL E$$ECTS ON PHOTOSYNTHESIS AND RESPIRATION Photosynthesis and photorespiration are sensitive to temperature changes, and respond directly to variations in temperature. 0igh temperatures favor o5ygenation over carbo5ylation. Carbo5ylation occurs during the dar6 phase, Calvin cycle, of photosynthesis and is the direct result of the activity of r%&"!#o. R%&"!#o a#t"va!e is an enzyme re.uired to change rubisco from the inactive to the active form in order to carry out carbo5ylation. "ubisco activity is sensitive to temperature/ The temperature-dependent association of rubisco activase with the thylakoid membrane was due to a conformational change in the rubisco activase itself, not to heat-induced alterations in the thylakoid membraneDuring a sudden and unexpected exposure of plants to heat stress, rubisco activase is likely to manifest a second role as a chaperone in association with thylakoid-bound ribosomes, possibly protecting, as a first aid, the thylakoid associated protein synthesis machinery against heat inactivation. R%&"!#o a#t"va!e: a e'()e *"t+ a te)perat%re,-epe-et -%a. /%#t"o0 Ro11a A, 2+a3 L, Aro EM. Plant 7. 899: 2eb-8+';*/;<=-7:. http///www.ncbi.nlm.nih.gov/entrez/.uery.fcgi>cmd#"etrieve?db#Pub$ed?list@uids#::8<9+98?dopt#1bstract he et p+oto!(t+et"# rate is the difference between the rate of carbon upta6e in photosynthesis and the rate of carbon lost in respiration. he net photosynthetic rate varies in plants depending on the environment in which the plant lives. Plants living in cooler climate have a lower ma5imum 'ma5* and minimum temperature 'mim* in which photosynthesis approaches zero. heir temperature optimum is lower, opt Aiochemical and physiological adaptations allow the plant to shift its optimum upta6e of carbon through photosynthesis toward the prevailing temperature of the environment. his process can also be observed during the seasonal shifts of temperature/ acclimation. P)P carbo5ylase is found in C1$ and C; plants, and is absent in C= plants. C= and C; show consistent differences in their photosynthetic response to temperature. here is no photorespiration during the initial carbon fi5ation by P)P carbo5ylase in C; plants. he opt of P)P carbo5ylase is higher than rubisco and high temperatures have little effect on C; plants. he opt for the C= pathway approaches that of rubisco. he opt for C; corresponds to the range of temperatures in which the activity of both enzymes, rubisco and P)P carbo5ylase, is relatively high. 1dditional information about the influence of temperature in the activity of rubisco/ http///www.ncbi.nlm.nih.gov/entrez/.uery.fcgi> cmd#"etrieve?db#Pub$ed?list@uids#::8<9+98?dopt#1bstract http///www.pnas.org/cgi/content/full/B7/8;/:8B=7 TEMPERATURE AND PLANT GRO4TH Plants re.uire certain amount of photosynthetic activity accumulated over time to reach certain point of development/ ma5imum growth, flowering, ripening of seeds, etc. Plants re.uire a number of degree-days of growth 'photosynthetic activity* to reach maturity or bloom.
he rates of photosynthesis vary with the time of the day and the season. Ct stops above and below certain temperatures, ma5 and min. he optimum temperature occurs between these two values. he inde5 of -e3ree,-a(! is used to relate growth to variations of temperature in a single season. he inde5 of degree-days is the sum of the departures in temperatures above some )"")%) or &a!e te)perat%re. he minimum temperature is selected as the temperature at which photosynthesis approaches zero. he mean daily temperature reflects growth and carbon accumulation. a6en from http///ohioline.osu.edu/agf-fact/9:9:.html O+"o U"ver!"t( E5te!"o, Depart)et o/ Hort"#%.t%re a- Crop S#"e#e. !ormula" #DD $ %T &igh plus T 'ow( divided by ), minus *+ The following ad,ustments are necessary" -( temperatures below *+ degrees ! are set at *+ degrees !, and )( temperatures above ./ degrees ! are set at ./ degrees !. This method of calculating #DD is often referred to as the %./,*+( system. 0xamples of #DD 1alculations" !or &igh $ .+ degrees !, 'ow $ /+ degrees !"#DD $ .+ plus /+ divided by ) minus *+ $ )+2 !or &igh $ /+ degrees !, 'ow $ 3+ degrees !"#DD $ /+ plus *+%3+( divided by ) minus *+ $ *2 !or &igh $ 4+ degrees !, 'ow $ 5+ degrees !"#DD $ ./%4+( plus 5+ divided by ) minus *+ $ ). #rowing Degree Days or heat units are calculated for each day starting the day after planting. Chec6 this site for a good e5planation of Ddegree-daysE/ +ttp:66***."p).%#-av"!.e-%64EATHER6--#o#ept!.+t). E7TREME TEMPERATURES AND PLANT SURVIVAL 2reezing temperatures can result in the formation of intra- and e5tracellular ice as well as phase change in membrane lipids. 2actors associated with the ability of cells to withstand freezing temperatures/ Cncreased solute concentrations. Fnsaturated lipids 'soluble fats* increase. Lipid concentration increases. 1mino acids are removed from proteins/ depolimerization. Cell membrane becomes more permeable. &mall cell size 1bscisic acid. 1bscisic acid accumulates in the leaves with dehydration. hrough its effects on second messengers such calcium ions, potassium ion channels open in the guard cells causing a massive lost of potassium. his loss of ions from the guard cells, causes water to leave the cells and the subse.uent loss of turgor of the guard cells closes the stomata. Critical minimum temperature/ 9G to :9GC and -:+G to -;9GC. Cn region where the temperature falls between -:+G to -;9GC, the dominant vegetation consists of broad- leaf deciduous plants. hese plants lower their freezing point by supercooling, the lowering of the freezing point by increasing solute concentration. Cells can lower the freezing point by no more than =GC by increasing solute concentrations. =9 genes were found in 'arix kaempferi that increase the supercooling capability to -<9HC. &ee abstract at http///I5b.o5fordIournals.org/content/+J/:=/=7=:.abstract>sid#<:b<;7e7-b;77-;b7 4ater in the cell wall and middle lamella freezes first with dropping temperature and releases heat 'heat of fusion or specific heat of melting*, which is absorbed by the adIacent cells and helps them to remain li.uid. hen, water moves out of the cells attracted to the ice crystals. olerance to freezing is not uniformly distributed through a plant. "oots, bulbs and rhizomes are the most sensitive to freezing '-:9 to -=9GC*. erminal buds are less resistant than lateral buds. 4oody stems are more resistant than buds and leaves. Changes in the ultrastructure of the cells have been observed/ multiple small vacuoles, increase in the number of vesicles, etc. 2or more details see/ http///aob.o5fordIournals.org/cgi/content/full/B9/+/<=7 6nnals of 7otany 89/ <=7-<;+, 8998, $arzanna &tefanows6a, $ieczys aw Kura and 1lina Kacpers6a 0airs insulate by trapping air and heat. he growth habit also helps in surviving freezing temperatures/ the interior temperature of cushion and rosette plants may be 89GC higher than the surrounding air. 1 high temperature of ;+GC disrupts metabolic processes. 0eat shoc6 proteins are involved but their role is not well understood. Cacti can maintain protein synthesis as fast as proteins brea6down and, in this way, avoid ammonia poisoning. $orphological and nactic movements allow plants to adIust to high temperatures, e.g. folding of leaves, spines, narrow leaves, photosynthesis carried out in the stem, changing the orientation of leaves to a parallel position to sun rays. PROCESSES OTHER THAN SURVIVAL AND GRO4TH emperature affects germination, reproduction, flower formation, flower unfolding, and ripening of fruits. emperatures between L= and %:=GC are needed by certain annuals and biennials to flower normally in the spring. PLANT RESPONSE TO 4ATER Mpen stomata allow CM8 into the leaves for photosynthesis, but also allow the water to escape, transpiration. 4ATER UPTA:E AND THE SOIL,PLANT,ATMOSPHERE CONTINUUM 1 water potential gradient e5ists between the soil, and the tissues of the roots, the stem and branches, the leaves, and the atmosphere. his water potential gradient is responsible for the movement of water from the soil through the plant into the atmosphere. he units used to describe the water potential are megapascals, $Pas. 4ater flows from areas of high water potential 'N* to areas of low water potential. his is called o!)ot"# potet"a.. he soil has the highest water potential and the atmosphere the lowest. Oatmosphere P Nleaf P Nstem P Nroots P Nsoil he movement of water across a membrane is called o!)o!"!. he osmotic potential of the cells 'concentration of solutes in the cytoplasm*, the matric potential or tendency of water molecules to adhere to soil particles, and the pull of gravity or gravitational potential, all influence the total water potential in the body of the plant. 1s plants lose water through transpiration, the solute in the cells becomes more concentrated, the water pressure drops in the cells, and water moves in from the areas of higher concentration. 1s water moves from the soil into the roots, the water potential of the soil drops and becomes more negative. he tendency of water to adhere to surfaces is called )atr"# potet"a.. 1s the water content of the soil drops, the remaining water adheres more tightly to soil particles and the matric potential drops. Cohesion between the water molecules also plays a role increasing surface tension in the soil pores between the clay particles and creating menisci 'sing. meniscus*. 1dhesion to clay particles and the formation of menisci can increase the matric potential significantly and ma6e it unavailable to plant roots. he te5ture of the soil affects the matric potential. Clay provides more surface than do sand and maintains a more negative matric potential. 1s the soil water potential drops, it becomes more difficult for the plant to maintain its water potential and eventually it cannot absorb more water. 1t this point, the stomata close to prevent the loss of water through transpiration but this also prevents the entry of CM8 into the leaves and disrupts photosynthesis. he value of the leaf water potential at which the stomata close varies with the species, and depends on the biochemistry, physiology and morphology of the species. 4ater must overcome the pull of gravity in order to move up the vascular tissue of the plant. he gravitational potential is a factor of the height of the plant from a reference point. Ct is positive above the reference point and negative below. Ct is important in the movement of water in tall trees. he gravitational potential, Ng, increases by 9.9: $Pa m -: above the ground. RESPONSE TO SHORT,TERM MOISTURE STRESS he closing of the stomata prevents the loss of heat by transpiration but the plant continues to intercept radiation and its internal temperature rises. 1n increase in internal temperature results in heat stress that interferes with protein synthesis and if prolonged, with chlorophyll synthesis. he plant may respond by curling its leaves, wilting, and dropping the leaves prematurely. he oldest leaves are shed first. Cf the drought continues, the tender twigs and branches die bac6. &ome plants under water-stress reduce their osmotic potential by accumulating ions of Ca 8% , $g 8% , K % and Qa 8% , and amino acids, sugars and sugar alcohols. he lower water potential of the leaves maintains the potential gradient from plant to soil. Conifers and evergreens may e5perience a browning and a diebac6 during the winter months due to water stress. Cf the temperature is high enough for water in the vascular tissue to li.uefy, the trees lose their water by transpiration but the water cannot be replaced because the ground is frozen. 3ehydration of the foliage occurs. PLANT RESPONSES TO LONG,TERM VARIATIONS IN 4ATER AVAILABILITY Cndividual plants growing under dry conditions have thic6er leaves than members of the same species growing under moist conditions. he leaves are thic6er because more layers of mesophyll are produced. here is more mesophyll per unit of area. $ore mesophyll layers increase photosynthesis but reduce the surface area that absorbs radiation and loses water through transpiration. "oot production increases under dry conditions by shifting the allocation of carbon from leaves to roots. Cndividuals growing under contrasting environmental condition show responses that compensate for the shortage of an essential resource. INTERPSECI$IC VARIATION IN ADAPTATIONS TO MESIC AND 7ERIC ENVIRONMENTS. Rerophytes have greater *ater %!e e//"#"e#( than mesophytes, that is, a greater rate of carbon upta6e per unit of water transpired. 4ater %!e e//"#"e#(/ rate of carbon upta6e per unit of water transpired. Photosynthesis/transpiration C; have higher water use efficiency than C= plants. C; plants maintain a very low concentration of CM8 within the mesophyll of the cells by having a great rate of carbo5ylation. his causes a steep gradient of CM8 concentration between the inside of the leaf and the outside air. he steeper CM8 gradient allows C; to maintain a higher rate of photosynthesis than C= plants for a given stomatal conductance. he ratio of root mass 'g* to leaf is 'cm 8 * increase with decreasing water availability. &pecies adapted to high and low resource availability show responses that compensate for the shortage of an essential resource/ Fnder 5eric condition, there is a lower stomatal conductance and lower rate of net photosynthesis than those species living in mesic environments. Lower stomatal conductance, however, results in greater water use efficiency. Lower allocation of carbon to production of leaves results in greater root production that increases the plant!s access to soil water. "educed leaf surface area and reduced photosynthesis results in less carbon upta6e and reduced growth. here is a trade off between higher rates of photosynthesis and growth when water is available and survival, growth and reproduction when water is consistently in short supply. LIN: BET4EEN PLANT 4ATER AND ENERGY BALANCE Plants dissipate heat through the loss of water by transpiration. Cf the water potential in the soil declines, the ability to absorb water also declines. Fnder mesic conditions, transpiration is the preferred means for heat dissipation. Plants also dissipate heat by convection. Cn 5eric environments, plants dissipate most its heat by convection. Leaf size decreases gradually as conditions change from mesic to 5eric. PLANTS RESPONSE TO $LOODING oo much water around the roots causes the death of root tips due to lac6 of o5ygen. "oot death follows due to poor absorption. 3etritus is added to the vascular tissue and the 5ylem clogs. 0igh water table causes plants to develop horizontal root systems that grow along the o5ygenated soil zone. PLANT ADAPTATIONS TO $LOODING Aere#+()a is a specialized tissue that contains air spaces that facilitate gas e5change and transport of air from shoots to roots. he porosity of plants adapted to flooding could be as great as <9,. Plants living in mesic or 5eric environments usually have a porosity of 8-7, space by volume. Pneumatophores are adaptations to permanently flooded environments. hey probably help in providing o5ygen to roots. 1erenchyma formation/ 4ithout o5ygen... "oots shift for aerobic to anaerobic respiration. Fpta6e of ions is inhibited he concentration o5ygen, potassium, nitrogen, and phosphorus decreases. )thylene is produced and accumulates. )thylene is insoluble and does not diffuse out of the roots and o5ygen upta6e is prevented. )thylene causes cells ne5t to the corte5 to brea6 and separate to form interconnected gas- filled chambers, aerenchyma tissue. &tem and lenticel +(pertrop+( H(pertrop+( is the enlargement of an organ without an increase in the number of constituent cells. 1n e5ample of this is &%ttre!!"3 or &%tt !*e.. which is an increase in the diameter at the base of the stem. he role of this seems to be to increase air space which allows for increased movement of gases. Aesides that, the wide base provides e5tra support for shallow rooted structures on a soggy substrate. PLANT ADAPTATIONS TO SALINITY &alts originate from the weathering of roc6s, irrigation and floods, human and animal additions and fertilizers. 1s salinity increases, plants have more difficulty in e5tracting water from the soil. Cn salty environments either soil or water, the water potential is lower than that of the cells or water tends to leave the cells. )ventually the cells dehydrate and plasmolyze. 0igh concentration of organic ions may be to5ic, e.g. high 1l =% is suspected to interfere with $g 8% upta6e. 8alt toxicity comprises osmotic and ionic components both of which can severely affect root and shoot growth. 9ptake of :a ; across the plasma membrane is very fast resulting in physiological effects on extracellular as well as intracellular sites. 8odium reduces binding of 1a ;; to the plasma membrane, inhibits influx while increasing efflux of 1a ;; , and depletes the internal stores of 1a ;; from endomembranes. These changes in the cell 1a ;; homeostasis are suggested here to be the primary responses to salt stress that are perceived by root cells. 8alt would almost instantly reduce the amount of 1a ;; being transferred to the leaf cells, with 1a ;; activity dropping and :a ; activity rising in the apoplasm of leaf cells. <lant, 1ell and 0nvironment ':BB8* :+, <8+-<=8. T+e ro.e o/ #a.#"%) " !a.t to5"#"t(. S. ")QT)L. Department of <lant 8cience, =aite 6gricultural >esearch ?nstitute, 9niversity of 6delaide, #len @smond, 86 *+/3,6ustralia. http///www.blac6well-synergy.com/doi/pdf/:9.::::/I.:=<+-=9;9.:BB8.tb9:99;.5Usearch#,88salt ,89to5icity,89plants,88 0alophytes are plants adapted to salty environments. &oils with more than 9.8, salt content are considered salty. hese soils are common in desert regions. :. he endodermis is the first effective barrier to too much salt in the environments of halophytes. heir corte5 contains a large amount of salt but their leaves have much less salt. 8. &ome plants have specialized organs to dispose of e5cess salt. hese plants do not have very effective barrier to salt absorption and the e5cess salt must be eliminated by other means. &alt-e5creting glands selectively remove slat from the vascular tissue of the leaves. Mther plants accumulate salt in the leaf tissues and then shed the leaves. 1 few halophytes are obligate and re.uire salt in their environment to grow best, e.g. mangroves grow best in low salinity- &alicornia grows best in moderate salinity and growth decreases in low and high salinity. &ome non-halophytes are resistant to salt spray but others are particularly sensitive. 2leshy leaves are often found in halophytes/ storage of water, e. g. &alicornia or pic6elweed. 1dditional information/ http///www.shef.ac.u6/aps/mbiolsci/Ieni/dissertation.pdf http///www.botgard.ucla.edu/html/botanyte5tboo6s/lifeforms/halophytes/fullte5tonly.html PLANTS AND NUTRIENTS 1ll plants re.uire at least :< nutrients for growth. &ome plants re.uire additional elements. $acronutrients are those elements that are needed in large amounts. here are B macronutrients/ C, 0, M, K, P, &, Q, Ca, and $g. C, 0, and M form the bul6 of the body of the plant and they are derived mostly from 08M and CM8. Qutrients are released in to the soil by weathering processes and absorbed by plant roots and incorporated into their tissues. here is a nutrient cycle from the soil to the plant and bac6 to the soil. NUTRIENT UPTA:E AND PLANT PROCESSES 1vailability of nutrients and demand affects the nutrient upta6e by plants. Qutrient upta6e is mediated by enzymes. 1s the concentration of nutrients increase, the absorption rate increases. )ventually the plant reaches a ma5imum upta6e rate and any further increase in concentration does not affect the upta6e rate. he $ichaelis-$enten e.uation relates these to factors/ V ; <V)a5 7 Ce5t=6:) > Ce5t= V # rate of nutrient upta6e Vma5 # saturation upta6e rate- all enzyme molecules are bound to substrate. Ce5t # e5ternal concentration Km # value of Ce5t at which V is half of Vma5- +9, of the active sites are bound to substrate. Qutrients are the substrate. 1t low concentrations of nutrients, the enzyme e5ists in an e.uilibrium between both the free form of the enzyme and the enzymeLsubstrate comple5- the nutrient substrate is the limiting factor- increasing nutrient concentration also increases enzyme substrate comple5 at the e5pense of the free enzyme, shifting the binding e.uilibrium to the right. &ince the rate of the reaction depends on the concentration enzyme-substrate comple5, the rate is sensitive to small changes in nutrient concentration. 0owever, at very high nutrient concentration, the enzyme is entirely saturated with substrate 'nutrient*, and e5ists only in the enzyme-substrate comple5 form. Fnder these conditions, the rate is insensitive to small changes in nutrient concentration. he nutrient concentrations of plant tissues have a direct relationship to 6ey processes related to plant growth, survival and reproduction. )5ample/ over +9, of the total nitrogen in leaf tissues affects photosynthesis, including the synthesis of rubisco and chlorophyll. PLANT ADAPTATIONS TO VARIATIONS IN NUTRIENT AVAILABILITY ?. Root 3ro*t+ ver!%! !+oot 3ro*t+ here is an increase allocation of carbon to root production with declining nutrient availability. he allocation of carbon to roots varies between species living in nutrient rich or nutrient poor habitats. &pecies living in nutrient rich habitats continue to increase its growth rate as nutrient availability increases. &pecies living in nutrient poor habitats declines after a short increase in growth as the nutrient concentration increases. &pecies living in a moderated nutrient environment increase its growth up to a point and reaches a plateau. Trowth response was measured as the accumulation of dry weight over the period of the e5periment. @. Lea/ .o3ev"t( )5periments have shown that species with short-lived leaves tend to have higher leaf nitrogen concentrations than those species having longer-lived leaves. &pecies with short-lived leaves tend to have a higher rate of photosynthesis than those with longer-lived leaves/ an inverse relationship between leaf life span and photosynthetic rate. here is a cost in producing a leaf in carbon and essential nutrients. Long-lived leaves are found in nutrient poor habitats, so when a leaf is produced it lasts long because there is slow nutrient upta6e due to the low availability of nutrients. Cf leaves are shed often in a low nutrient environment, the plant will lose more nutrients than it can ta6e from the soil resulting in a net loss and eventually in death. A. I/.%e#e o %tr"et ava".a&"."t( "oots ta6e up nutrients in soil solution as water is absorbed. 1ctive transport of nutrient also occurs. 1s nutrients are ta6en from the soil, a zone of nutrient depletion is formed around the roots. Qutrients flow into this zone of nutrient depletion from the surrounding areas as a result of the diffusion gradient established by the root upta6e. 1s leaves become old, senescent, nutrients are removed and transported to perennial part of the plant to be reused. his process is called %tr"et retra!.o#at"o. M%t%a."!t"# re.at"o!+"p! between plants and mycorrhizae and nitrogen-fi5ing bacteria increase nutrient availability to plants. "hizobium bacteria are the nitrogen-fi5ing bacteria associated with the roots of plants. "hizobium depends on the carbon provided by the plant as a source of energy, and in turn they provide the plant with nitrogen. Cyanobacteria, blue-green algae, are the nitrogen-fi5ing organisms found in a.uatic environments. hese mutualistic associations are most beneficial in environments with low-nutrient availability. Cn environment with high nutrient availability it represents a cost since the plant must provide photosynthates to support the bacteria and mycorrhizal fungi with little benefit. CALCICOLES AND CALCI$UGES &oil acidity affect nutrient upta6e because acidity affects the solubility of minerals. Ca.#"#o.e plants are those that prefer a basic or al6aline soil- soils rich in calcium slats. Ca.#"/%3e plants are those that prefer an acid soil. Calcifuge means lime-hating- low in calcium salts. Ne%trop+".%! plants are those that tolerate either condition. Low p0 is associated to calcium deficiency. 0ighly acidic soils contain high amounts of aluminum and iron, which are to5ic to many plants. 2ree aluminum accumulates on the surface of the root and in the root corte5. it interacts with phosphorus to form highly insoluble compounds. PLANTS O$ SERPENTINE AND TO7IC SOILS. &erpentine is a magnesium-iron silicate, '$g, 2e*8&iM;. Ct also contains Ca, 1l, Qa and i. Ct may contain chromite, 2eCr8M; and garnierite, '$g, Qi*&iM= W n08M &i, $g and 2e are the maIor constituents- 1l is low. Concentration of Ca and heavy metals is low but plays an important role in the soil. 0eavy metals such as iron, nic6el, chromium, cesium, zinc and cobalt are to5ic to plants, causing chlorosis and stunted growth. hey interfere with nutrient upta6e and root growth and penetration. 0igh concentrations of heavy metals 'Qi, Cr, Co*, and $g, a low Ca/$g ratio and low fertility characterize serpentine soils- they are low in Ca, P, Qa, and 1l. &erpentine soils contain a flora tolerant of these conditions. A<lants can be strikingly different on serpentine soils for several reasons. Binerals that contain high levels of nickel and chromium are relatively common in serpentine soils and can cause toxicity in plants. 6lthough some serpentine soils can be deep, most are shallow, restricting water holding capacity and rooting depths. 6dditionally, serpentine soils have nutrient deficiencies and imbalances. ?mportant nutrients like potassium, phosphorous, nitrogen, and molybdenum have been found to be in very low amounts in most serpentine soils. @verall, the most consistent restriction for plant growth appears to be the extraordinary low level of calcium compared to magnesium. 6lthough both are necessary macronutrientsCand plants can be selective in absorption of specific mineralsCcalcium and magnesium in plant available forms are similar chemically and can compete for adsorption. 1alcium deficiencies result. 8ome serpentine endemics have adapted by having an ability to accumulate calcium and exclude excess magnesium. @thers isolate and store toxic heavy metals.A 2ran6 "auchschwalbe, ;/=9/9;. http///w.w.w.ucce.ucdavis.edu/counties/cetuolumne/ &erpentine soils have a high proportion of endemic and ecotype species. E-e)"#! are species restricted to a particular habitat or geographical region. E#ot(pe! are ecological races well adapted to a local set of conditions hese types tolerate Qi and Cr in their tissues at levels highly to5ic to other plants. &ome plants have developed mechanisms that e5clude heavy metals from the plant. 1 few species grow only in the presence of certain heavy metals and are indicators of the presence of those metals in the soil. he selective influence of heavy metals... 2avors tolerant seedlings from the surrounding area. Continue selection against susceptible genotypes despite gene flow from the surrounding population. &election for the ability to survive in physically harsh, largely 5eric, nutrient-poor environment.