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Notice: ©1990 Springer. This manuscript is an author version with the final publication available at http://www.springerlink.com and may be cited as: McClintock, J. B., Cameron, J. L., & Young, C. M. (1990). Biochemical and energetic composition of bathyal echinoids and an asteroid, holothuroid and from the Bahamas. Marine Biology, 105(2), 175‐183. doi:10.1007/BF01344284

('!J' 1) Marine Biology 105. 175-183 (1990) Marine ==: Biology © Springer-Verlag 1990

Biochemical and energetic composition of bathyal echinoids and an asteroid, holothuroid and crinoid from the Bahamas

J. B. McClintock 1, J. L. Cameron 2 and C. M. Young 2

1 Department of Biology. University of Alabama at Birmingham. University Station, Birmingham. Alabama 35294, USA 2 Department of Larval Ecology, Harbor Branch Oceanographic Institution. 5600 Old Dixie Highway. Fort Pierce. Florida 34946. USA

Date of final manuscript acceptance: February 2, 1990. Communicated by 1. M. Lawrence. Tampa

Abstract. The biochemical 'and energetic composition of Walker et al. (1987 a, b) reported on the biochemical and body components often species ofbathyal echinoids, and energy content ofcommon bathyal and abyssal elaspodid an asteroid, a holothuroid and a stalked crinoid were and aspidochirote holothurians from the Atlantic Ocean. determined from individuals sampled from a variety of These authors noted that dense populations of echino­ deep-water sites near the Bahamas (north Caribbean Sea) derms often dominate megafaunal biomass in deep At­ in October 1988. When compared with other studies of lantic Ocean waters. At bathyal depths (300 to 900 m) in , no geographic- or depth-related differences the vicinity of the Bahamas in the northern Caribbean in biochemical or energetic composition were found. Sea, benthic megafaunal biomass is also dominated by Body-wall tissues were composed primarily of skeletal echinoderms (1. Miller personal communication). Regu­ material (mineral ash), but were comparatively high in lar and irregular echinoids are particularly abundant. organic material in the echinothuriid echinoids, and the and holothuroids and stalked occur in large pop­ asteroid and holothuroid. Gut tissues and pyloric cecae ulations (c. Young and L. Cameron unpublished data). had high levels of lipid and protein, indicating their po­ The purpose of the present study was to provide in­ tential role in nutrient storage. Body-wall tissues were formation on the biochemical and energetic composition generally low in energy, but were highest in the echinoids ofsomatic tissues of bathyal echinoderms. This informa­ belli (7.7 kJ g " I dry wt) and Sperosoma antil­ tion is of comparative interest with regard to shallow­ lense (8.0 kJ g- I dry wt), the asteroid Ophidiaster alexan­ water species and species from temperate and polar lati­ dri (8.9 kJ g " I dry wt), and the holothuroid Eostichopus tudes. and may provide insight into the functional role of regalis (13.1 kJ g - I dry wt). Energy levels of gut and body-tissues. Moreover. when coupled with pyloric cecal tissues were two to three times higher than measurements of echinoderm densities. these data can be those of body-wall tissues. Total somatic tissue energy used to estimate the energetic densities of bathyal echino­ values varied greatly among species, ranging from 1.5 kJ derm populations. in the echinoid Aspidodiadema jacobyi to 142.1 kJ in E. regalis. As the bathyal echinoderms examined in this study occur in great abundance. they represent a signifi­ Materials and methods cant reservoir of organic and inorganic materials and Adults of ten species of echinoids. the asteroid Ophidiastcr alexan­ energy in deep-water benthic systems. dri. the holothuroid Eostichopus regalis, and the crinoid decorus were collected using a manned submersible (Table 1). The location and depth of collection was recorded for each species. Individuals were dissected aboard ship. and the somatic tissues were Introduction frozen until lyophilization. Echinoids were dissected into body wall, lantern. and gut tissues. In species with long spines. the spines were separated from the body wall and treated as a separate body compo­ The biochemical and energetic cornposruon of echino­ nent. The asteroid was dissected into body wall and pyloric cecae. derm body-components has been reported for a number the holothuroid into body wall and gut. and the crinoid into arms. of shallow-water tropical, temperate, and polar environ­ stalks. and cirri. Intact body components were weighed. subsarn­ ments (Giese 1966. 1976. Lawrence 1973, Lawrence and pled. lyophilized. reweighed. and ground into a line powder in a Guille 1982. Magniez 1983, McClintock and Pearse Wiley Mill. 1987 a, b, Wacasey and Atkinson 1987. McClintock TCA-soluble carbohydrate and NaOH-soluble protein were measured spectrophotornetrically using the techniques of Lowry 1989 a, b). However. few studies have examined the bio­ et al. (1951) and Dubois et al. (1956). respectively. Chloroform­ chemical and energetic composition of bathyal and methanol-soluble lipid was determined gravimetrically using the tech­ abyssal echinoderms. Sibuet and Lawrence (1981) and nique of Freeman et al. (1957). Ash was measured by placing lyo- 176 J B. McClintock et al.: Biochemical/energetic composition of bathyal echinoderms

Table 1. Collection locations and depths for bathyal echinoderms collected in October 1988 for biochemical and energetic analyses

Species Longitude; Location Depth Latitude (m)

Class Echinoidea Subclass Order Family 23Q55.9'N; French Bay, 484-493 74°33.52'W San Salvador Styiocydaris lineata 24°2.25'N; Tartar Bank, 617 75Q28.15'W Cat Island

Subclass Superorder Diadematacea Order Family Araeosoma belli 24D2.01'N; Cockburn Town. 676-722 74°32.40'W San Salvador 24' 3.55'N; Cockburn Town, 676-722 74°32.29'W San Salvador Sperosoma antillense 24°6.35'N; Rocky Point 291 74°32.7'W San Salvador

Order Family Aspidodiadema jacobyi 24°2.01'N; Cockburn Town, 728 74"32.40'W San Salvador

Superorder Order Family Archaeopneustes histrix 24'2.01'N; Cockburn Town. 607 74'32.40'W San Salvador Linopneustes longispinus 24'02.5'N: Cockburn Town, 662 74°32.40'\\, San Salvador Palaeopneustes cristatus 2402.5'N: Cockburn Town. 584 74'32.40'W San Salvador Palaeobrissus hilgardi 24'02.5N; Cockburn Town, 637-659 7432.4'\\, San Salvador

Class Stelleroidea Subclass Asteroidea Order Valvatida Family Ophidiasteridae Ophidiastcr alexandri 24 2.25'N; Tartar Bank. 636-669 752lU5'W Cat Island

Class Holothuroidea Order Aspidochirota Family Stichopodidae Eostichopus regalis 25 27.52'N: Egg Reef 421 7654.32'\\,

Class Crinoidea Order Family 23 57.42'N: Sandy Point. 479 7434.45'\\' San Salvador

philized tissues into a muffle furnace at 500 'C for 4 h (Paine 1971). The total amount of energy allocated to each somatic tissue was The remaining organic refractory material was considered to be calculated by multiplying the kJ g I dry wt tissue of each compo­ NaOH-insoluble protein (Lawrence and Kafri 1979). The energetic nent by the total dry wt of [he intact body component. These values composition of the tissue was calculated by multiplying the values were summed, yielding the total somatic energy content ofan indivi­ of each of the organic classes by energy equivalents (Brody 1945) dual. IS 1.B. McClintock et al.: Biochemical/energetic composition of bathyal echinoderms 177 An ANOYA was used to make comparisons of body-wall bio­ Results chemical compositions within those groups of echinoids whose body-wall tissues were combined with short spines, and those whose The biochemical composition ofthe body-wall tissues are body-wall tissues were analyzed separately. Percentage data were transformed before statistical analysis using an arc-sine transforma­ presented in Table 2. The body-wall tissues of echinoids tion. When significant differences were detected with the ANOYA, from which the body wall and spine measurements were 95% confidence limits allowed an evaluation of where significant combined were composed primarily of mineral ash, with differences occurred. values ranging from 69.8% dry wt in the echinothuriid

100 Ash F - 168.44. P = 0.000 100 Ash F ~ 100.05. P = 0.000 ~o 90 - - 80

80 n .. 1.0 Soluble Carbohydrate F = 0.47, P - 0.755 1.0 - Soluble Carbohydrate F = 4.08. P = 0.044 0.8 0.8 -t- 0.6 + - - r+ 0.6 ~ 0.4 r- - 0.4 0.2 0.2 0.0 r:1 e- I o 0.0 1 I 1 I ...­ c ....­ CIl 10.0 Soluble Lipid c (J F = 2.72, p ~ 0.080 CIl 10.0 Soluble Lipid .. (J r ~ 1.58, P = 0.261 CIl .. o, 8.0 CIl a, 8.0 6.0 6.0 4.0 4.0 2.0 2.0 0.0 O.a-t--...... ,.'"'------'Y--...... ,.'"'------'Y----,

20.0 Soluble Protein F - 31.43. p = 0.000 20.0 Soluble Protein r = 19.12. P = 0.000 15.0 r+ .-- r- 15.0 10.0 10.0 5.0 5.0 0.0 n n I I o. a+--....,..L---'-,r'-----J.,..L---Y---, ..s> i;:, ~ i)- 0 6 -s ~ ~ 0 -o t' :.§' 0 6' ~ ii' .~ f ~ ::s e o~ s .::;," J5 e 0 .r;> ~ .;; t ...... ~ E -0 et' 0 0" ~ ,f ~ ~ 0 ,?Q. Q..0 Q..-<: Fig. 1. Biochemical composition (ash. soluble carbohydrate. solu­ Fig. 2. Biochemical composition (ash. soluble carbohydrate. solu­ ble lipid. soluble protein) of body wall and associated short spines ble lipid. soluble protein) of body wall (no spines included) of four of five species of bathyal echinoids collected in October 1988 from species of bathyal echinoids collected in October 1988 from Ba­ Bahamas. Histograms show means (± 1 SE: bars) of three individ­ hamas. Histograms show means (± 1 SE; bars) of three individuals uals except for P. hilgardi (n = 5). Also shown are F values and except for A. histrix (n=4). Also shown are Fvalues and associated associated probability estimations for each ANOYA probability estimations for each ANOYA 178 1. B. McClintock et al.: Biochemical/energetic composition of bathyal echinoderms

Table2. Biochemical composition (% dry wt ) of bathyal echinoderms collected from Bahamas in October 1988. Values are means for sample sizes of 2; means ± 1 SD are given for sample sizes> 2.....: 2 and 3 pooled samples. respectively

Species n Ash Carbohydrate. Lipid. Protein soluble soluble soluble insoluble

Class Echinoidea Calocidaris micans body wall 3 90.0± 1.4 O.HO.1 2.7± 1.7 3.3± 1.4 3.5 ± 1.4 spines 1 93.7 0 0.4 2.0 3.9 lantern 1·· 59.0 0.6 1.2 4.0 35.2 gut 1·· 15.5 1.8 17.9 28.8 36.0 lineata body wall 3 90.2±0.6 0.6±0.3 3.8±0.8 3.9±0.8 1.4±0.8 spines 1 96.6 0.4 1.2 0 1.8 lantern 3 89.4 ± 1.3 0.5±O.1 1.8 ± 2.0 5.1 ±3.1 3.2±0.1 gut·· 1 25.3 1.2 16.5 36.2 20.7 A raeosoma belli body wall +spines 3 70.7±2.3 0.5±0.4 3.9±2.5 14.7 ± 5.9 11.4 ± 2.4 lantern 3 82.3 ±2.4 0.HO.5 2.7±1.0 5.2±3.4 9.3±3.2 gut 1·· 20.5 1.8 21.5 32.1 24.1 Phormosoma placenta body wall +spines 3 76.01 ± 1.7 0.6±0.2 3.8±1.1 13.9±3.7 5.7±2.4 lantern 3 83.5±2.3 0.5±0.1 1.8±0.1 6.6±2.4 7.4±3.9 gut 1 43.9 1.4 141 27.2 13.4 Sperosoma antil/ense body wall + spines 3 69.8 ± 2.6 0.6±0.1 6.1 ±1.6 14.1 ±2.1 9.4±2.4 lantern 3 84.5 ± 1.6 0.5±0.1 3.0± 1.5 8.6± 1.1 3.3±2.0 gut 1 23.2 1.6 21.6 30.4 22.2 Aspidodiadema jacobyi body wall 3 74.0 ± 1.7 0.8±0.3 46±3.1 10.6±2.2 9.9± 1.2 spmes 1 86.3 1.1 1.8 5.6 5.2 lantern 1• 88.9 0.6 5.1 3.5 1.9 Archaeopneustes histrix body wall 4 88.4 ± 1.2 0.7±01 4.9±0.5 3.6±0.8 2.4±12 spines 1 91.8 0.7 0.2 0.5 7.2 gut 1• 35.3 0.9 10.7 29.5 23.6 Linopneustcs longispinus body wall 3 91.0±0.1 0.5±0.1 2.4±09 3.7±0.6 2.2 ± 1.2 spines 1 92.3 0.4 0.8 1.7 4.8 gut 1·· 44.9 1.7 15.1 15.5 21.8 Pataeopneustcs cristatus body wall + spines 2 90.9 0.9 1.9 1.4 5.1 gut 1• 47.4 1.2 10.4 37.0 4.0 testes 1 19.2 07 32.9 20.4 26.8 Palacobrissus hilgardi body wall + spines 5 91.8 ± 1.0 0.5 ±0.2 2.6±06 1.6±0.9 3.5 ± 1.8 gut I •• 520 1.4 4.0 33.4 9.2

Class Stelleroidea Ophidiaster alcxandri body wall 64.6 0.7 40 19.2 115 pyloric ceca 112 1.7 23.2 32.3 31.6

Class Holothuroidea Eostichopus regalis body wall 2 51.6 1.4 10.7 18.4 18.0 gut 1 • 60.3 0.4 9.1 11.7 18.5 Class Crinoidea Neocrinus decorus arms 2 82.1 0.9 2.7 8.8 57 stalk 2 91.8 0.7 1.6 0.7 53 cirri 2 93.6 0.3 1.0 15 3.7 ps

J. B. McClintock et al.: Biochemical/energetic composition of bathyal echinoderms 179

Tabid. Energetic composition (kJ g " Idry wt) ofbathyal echinoderms collected from Bahamas in October 1988. Values are means for sample sizes of 2; means ± 1 SD are given for samples> 2. ". **: 2 and 3 pooled samples. respectively

Species n Carbohydrate. Lipid. Protein Total soluble soluble soluble insoluble

Class Echinoidea Calocidaris micans body wall 3 0.1 ±0.02 0.6±0.1 0.5±0.4 0.6±0.4 1.8 spines 1 0 0.1 0.5 0.9 1.5 lantern 1** 0.1 0.5 0.9 8.3 9.8 gut 1 ** 0.3 7.1 6.8 8.5 22.7 Stylocidaris lineata body wall 3 0.1 ±0.05 1.5±0.3 0.9±0.2 0.3±0.2 2.8 spines 1 0.1 0.4 0 0.4 0.9 lantern 3 0.1 ±0.01 0.7±0.2 1.2±0.7 0.7±0.1 3.6 gut 1** 0.2 6.6 8.6 4.9 20.3 Araeosoma belli body wall + spines 3 0.1 ±0.1 1.5± 1.1 3.5± 1.4 2.6±2.2 7.7 lantern 3 0.1 ±0.1 1.1±0.4 2.7±1.0 2.2 ± 1.0 6.1 gut 1 ** 0.3 8.5 7.6 5.7 22.1 Phormosoma placenta body wall + spines 3 0.1 ±0.02 1.6± 0.5 3.4±0.2 1.4±0.6 6.5 lantern 3 0.1 ±0.05 0.7±0.1 1.7 ±0.5 1.7±0.9 3.5 gut 1 0.3 5.5 6.4 3.3 15.5 Sperosoma antillense body wall + spines 3 0.1 ±0.01 2.4±0.6 3.3 ±0.5 2.2±0.6 8.0 lantern 3 0.1 ±0.02 1.2±0.6 2.0±0.3 0.8±0.5 4.1 gut 1 0.3 89 7.2 5.3 21.7 Aspidodiadema jacobvi body wall 3 0.1 ±0.05 1.9 ± 1.2 2.5±0.6 2.3±0.2 6.8 spines 1 0.1 0.7 1.4 1.2 3.4 lantern 1* 0.1 2.0 0.8 0.5 3.4 Archaeopneustcs histrix body wall 4 0.110.01 2.0±0.2 0.9±0.2 0.4±0.3 3.4 spines 1 0.1 0.2 0.6 1.7 2.0 gut 1* 0.2 4.2 6.9 5.6 16.9 Linopncustcs longispinus body wall 3 01 ±0.02 2.4 ± 0.6 0.9±01 0.HO.3 3.9 spines 1 0.1 0.3 0.4 1.1 1.9 gut 1** 0.3 6.0 5.4 3.7 15.4 Palaeopncustes crist at us body wall + spines 2 0.2 0.7 0.3 1.7 2.9 gut 1* 0.2 4.1 8.7 1.0 14.0 Palaeobrissus hilgardi body wall + spines 5 0.1 ±0.02 1.4±1.:' 0.3 ±0.2 0.7±02 2.5 gut 1** 0.3 1.5 78 ., ., 14.3

Class Stelleroidca Ophidiaster alcxandri body wall 0.1 1.6 4.5 2.7 8.9 pyloric ceca 0.3 9.2 7.6 7.5 24.6

Class Holothuroidea Eostichopus rcgalis body wall 2 0.2 4.3 4.4 4.2 13.1 gut 1* 0.1 3.6 2.7 4.3 10.7

Class Crinoidea Neocrinus decorus arms 0.1 1.0 2.1 1.4 4.6 stalk 0.1 0.6 0.2 1.2 2.1 CIrrI 01 0.4 0.3 0.9 1.7 .... 180 1.B. McClintock et aI.: Biochemical.energetic composition of bathyal echinoderms

Table 5. Length, height. dry wt, and kJ of individual somatic body components, and total somatic energy content of representative individuals of bathyal asteroid, holothuruid and crinoid collected 3 from Bahamas. Values in parentheses are percentages of total; Nil 6 not applicable

Parameter Ophidiaster Eostichopus Neocrinu- alcxandri regalli decorus (asteroid) (holothuroid) (crinoid)

Arm length (em) 6.2 Body length (em) 19.5 Body height (em) 72.5 Gravimetric composition (g dry wt) body wall 4.18 1007 arms 7.24 pyloric ceca 0.77 gut 0.95 stalk 907 cirri 1.76 Energetic composition (kJ) body wall 37.2 (66) 131.9 (93) arms 33.3 (60) pyloric ceca 18.9 (34) 19.0 (34) gut 10.2 (7) 3.0 (6) ,-- ,-- ,-- r- \0 r- stalk ~~~ cirri r--~v ~ 0001 Total 56.1 142.1 55.3

echinoid Sperosoma antillense to 91.8% dry wt in the spatangoid echinoid Palaeopneustes hilgardi. Levels of ash were significantly (P < 0.05) higher in the body wall and spines of S. antillense than In the other two

*re:, 'D 0 echinothuriids (Araeosoma belli, Phormosoma placenta), NN 100 while all echinothuriids had significantly (P < 0.05) lower ash levels than spatangoids (Fig. 1). The body-wall and spines of the spatangoid echinoids tStvlocidaris lineata

,--~ and Palaeopneustes hilgardi had significantly (P < 0.05) ~~ lower levels of NaOH-soluble protein than that of the c o \0 0' Cl echinothuriids (Fig. 1). No significant differences in lev­ r---: N-= """oc els oflipid or TCA-soluble carbohydrate were detected in the five echinoids for which body-wall and spines were combined. Body-wall tissues of echinoids for which only the body wall was examined also showed differences in V)c~oc - C 'D c '-0 '-D r--, \0 proximate composition. The small diadematoid echinoid "l oor-r------"'-/0 -,.,.. t- stes longispinus) (Fig. 2). ", ~ ~ components (arms, stalk. cirri: 82.1 to 93.6% dry wt). EC][ c :; Among the organic constituents of echinoid. asteroid. . ;; "'0.0 r.n ~ O(j ~~ and holothuroid body-wall tissues. levels of NaOH-solu­ o ble protein (1.4 to 19.2% dry wt) and insoluble protein ~------1.B. McClintock et al.: Biochemical/energetic composition of bathyal echinoderms 181 (refractory organic material: 1.4 to 18.0% dry wt) were skeletal material in the body-wall tissues of spatangoids generally highest. This w~s also true fo.r the arms, stalk. than cidaroids, echinothuriids and diadernatids may be and cirri of N. decorus. WIth the exception of the regular related to a decreased need for structural proteinaceous echinoid Sperosoma antillense and the holothuroid E. re­ material (connective tissue) to support the ossicles and galis, levels of lipid in body-wall tissues were generally plates of the test of the spatangoids. Levels of protein very low (1.9 to 4.6% dry wt). This was also the case for (NaOH-soluble and insoluble) were lower in the body­ the arms, stalk, and cirri of N. decorus. TCA-soluble car­ wall tissues of the spatangoids than the other echinoids bohydrate levels were very low in echinoid, holothuroid, examined. Echinoid spines were also high in mineral ash; and asteroid body-wall and all crinoid tissues (0.5 to small amounts of lipid and protein occur in species with 1.4% dry wt). an epithelial layer surrounding the spines. The lanterns of The organic composition ofthe gut tissues of the echi­ all but the spatangoids were generally high in mineral noids and holothuroid was dominated by NaOH-soluble ash, as found by Lawrence and Guille (1982) for shallow­ and insoluble protein (cumulative vales ranged from 30.2 water echinoids. However, the lantern of Calocidaris mi­ to 64.8% dry wt), with widely variable levels oflipid (4.0 cans had comparatively high levels of insoluble protein. to 22.6% dry wt), and low levels of TCA-soluble carbo­ Presumably this protein is structural in nature and a con­ hydrate (0.9 to 1.8% dry wt) (Table 2). stituent of connective tissue. Large amounts of connec­ The energy levels associated with each of the organic tive tissue associated with the lantern may reflect feeding constituents and energy levels (kJ g - I dry wt) of body habit, and the need to strengthen a lantern used to rasp tissues are presented in Table 3. Patterns of energetic at hard materials. composition were similar to biochemical composition. In The body-wall tissues of the asteroid Ophidiaster the echinoids, body-wall tissues were generally low in alexandri and the holothuroid Eostichopus regalis had energy (1.8 to 8.0 kJ g-I dry wt). Crinoid body compo­ higher organic contents and lower mineral ash content nents and echinoid spines and lanterns also had low levels than echinoids (however the body wall of the ofenergy. Higher levels ofenergy occurred in gut tissues, echinothuriids has a high organic content). This is related ranging from 10.7 kJ g - 1 dry wt in the holothuroid Eosti­ to a decrease in the degree of skeletalization and high chopus regalis to 22.7 kJ g - I dry wt in the echinoid levels of protein. Krishnan (1968) found that protein is Calocidaris micans. the primary organic constituent of body wall tissues of Total somatic energy contents of bathyal echino­ Holothuria scabra, and serves as a principal component derms (Tables 4 and 5) varied from a low of 1.5 kJ in the of the abundant connective tissues. These body-wall tis­ echinoid Aspidodiademajacobyi (test diameter = 1.4 em) sues may provide a substantial reservoir of nutrients and to a high of 142.1 kJ in the holothuroid Eostichopus re­ energy (Giese 1976, Prim et al. 1976, McClintock 1989 b, galis (body length = 19.5 em). Saito and Watts 1989). The ability to store nutrient re­ serves may be ofparticular importance in species occupy­ ing bathyal and abyssal depths. where nutrient availabil­ Discussion ity may be highly episodic (Sanders et al. 1965. Menzies et al. 1973. Rowe and Staresinic 1979). The arms and cirri The biochemical composition of the body components of of the bathyal stalked crinoid Neocrinus decorus are these bathyal echinoderms are similar to those from higher in skeletal material than the antarctic comatulid shallow-water temperate (Giese 1966, 1976, Lawrence Promachocrinus kerguelensis (McClintock and Pearse and Guille 1982, McClintock 1989 b), tropical (Lawrence 1987 a). As a supporting structure, it is composed primar­ 1973, Lawrence and Guille 1982) and polar (Lawrence ily of skeletal material. and Guille 1982. McClintock and Pearse 1987 a, b, Me­ The echinoid and holothuroid gut tissues contain 2 to Clintock et al. 1988) environments. Comparisons with 3 times higher levels of organic constituents than body­ other bathyal and abyssal echinoderms are restricted to wall tissues. Gut lipid levels as high as 10 to 23% dry wt holothurians (Sibuet and Lawrence 1981, Walker et al. were common in bathyal echinoderms. High species­ 1987 a. b), of which only one species was investigated in specific variations in levels oforganic constituents reflect the present study. The biochemical composition of the the dynamic nature of this tissue and its possible role in body wall and gut tissues of the bathyal holothuroid nutrient storage. Klinger et al. (1988) found that the gut Eostichopus regalis was similar to the 17 species of tissues of the echinoid Lytechinus variegatus may serve bathyal and abyssal holothuroids examined in earlier for short-term nutrient storage. The pyloric caeca of the studies. Although the small number of asteroids, asteroid Ophidiaster alexandri were also high in organic holothuroids, and crinoids examined precludes a broad constituents and could have a nutrient-storage function. comparison among these groups. clearly the differences The energetic level of the body-wall tissues of echi­ in biochemical composition of bathyal echinoids cannot noids and the arms, stalk. and cirri of the crinoid were be related to geographic or depth-specific factors. low (1.8 to 4.6 kJ g - 1 dry wt). However. the echinoids Lawrence and Guille (1982) indicated that the similarity Araeosoma belli, Phormosoma placenta, Sperosoma antil­ in composition reflects fundamental characteristics of lense and Aspidodiadema jacobyi had body-wall tissues echinoderm body-components which determine their with twice these levels ofenergy (6.5 to 8.0 kJ g - I dry wt). composition. Higher levels of energy are attributable to energy bound The body-wall tissues of bathyal echinoids are com­ in soluble and insoluble protein. and may reflect greater posed primarily of skeletal material. The greater levels of amounts ofconnective tissue and lesser amounts of skele- <

182 1.B. McClintock et al.: Biochemical/energetic composition of bathyal echinoderms

tal elements in the body wall of these echinoids. Araeoso­ Giese. A. C. (1966). On the biochemical constitution of some rna belli possesses a flexible test. Apparently its vulnera­ echinoderms. In: Boolootian, R. A. (ed.) Physiology of Echino­ bility to predation is offset by the presence of poisonous dermata. Wiley. New York, p. 757- 796 Giese. A. C. (1976). Physiology of the echinoderm body wall. Tha­ spines. The energy content of the body-wall tissues ofthe lassia jugosl. 12:153-163 asteroid Ophidiaster alexandri was similar to these echi­ Khripounoff, A.. Sibuet, M. (1980). La nutrition dechinoderm-, noids, while the body-wall tissues of the holothuroid abyssaux. I. Alimentation des holothuries. Mar. Biol. 60: 17-26 Eostichopus regalis contained higher levels of energy Klinger, T S., Watts. S. A., Forcucci, D. (1988). Effect ofshort-term (13.1 kJ g-I dry wt). This is the result of high levels of feeding and starvation on storage and synthetic capacities of gut tissues of Lytechinus variegatus (Lamarck) (Echinodermata: energy derived from protein. Walker et al. (1987 a, b) also Echinoidea). 1. expo mar, BioI. Ecol. 117: 187-195 found high levels ofenergy associated with protein in the Krishnan. S. (1968). Histochemical studies on reproductive and body wall of bathyal and abyssal holothuroids. nutritional cycles of the holothurian, Holothuria scabra. Mar. The bathyal echinoderms examined in this study are Biol. 2: 54-64 very patchy in their distribution but often occur in high Lawrence, 1. M. (1973). Level, content and caloric equivalents of the densities and are the most abundant benthic megafauna lipid. carbohydrate, and protein in the body components of (c. Young and L. Cameron personal observation). 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