char = P;code = 80 char = e;code = 101 char = e;code = 101 char = l;code = 108 char = i;code = 105 char = n;code = 110 char = g;code = 103 char = ;code = 32 char = t;code = 116 char = h;code = 104 char = e;code = 101 char = ;code = 32 char = o;code = 111 char = n;code = 110 char = i;code = 105 char = o;code = 111 char = n;code = 110 char = :;code = 58 char = ;code = 32 char = r;code = 114 char = i;code = 105 char = b;code = 98 char = o;code = 111 char = s;code = 115 char = o;code = 111 char = m;code = 109 char = e;code = 101 char = s;code = 115 char = ;code = 32 char = a;code = 97 char = r;code = 114 char = e;code = 101 char = ;code = 32 char = a;code = 97 char = n;code = 110 char = c;code = 99 char = i;code = 105 char = e;code = 101 char = n;code = 110 char = t;code = 116 char = ;code = 32 char = m;code = 109 char = o;code = 111 char = l;code = 108 char = e;code = 101 char = c;code = 99 char = u;code = 117 char = l;code = 108 char = a;code = 97 char = r;code = 114 char = ;code = 32 char = f;code = 102 char = o;code = 111 char = s;code = 115 char = s;code = 115 char = i;code = 105 char = l;code = 108 char = s;code = 115 char = .;code = 46 result sql = Peeling the onion: ribosomes are ancient molecular fossils.
Peeling the onion: ribosomes are ancient molecular fossils.char = W;code = 87 char = e;code = 101 char = ;code = 32 char = d;code = 100 char = e;code = 101 char = s;code = 115 char = c;code = 99 char = r;code = 114 char = i;code = 105 char = b;code = 98 char = e;code = 101 char = ;code = 32 char = a;code = 97 char = ;code = 32 char = m;code = 109 char = e;code = 101 char = t;code = 116 char = h;code = 104 char = o;code = 111 char = d;code = 100 char = ;code = 32 char = t;code = 116 char = o;code = 111 char = ;code = 32 char = e;code = 101 char = s;code = 115 char = t;code = 116 char = a;code = 97 char = b;code = 98 char = l;code = 108 char = i;code = 105 char = s;code = 115 char = h;code = 104 char = ;code = 32 char = c;code = 99 char = h;code = 104 char = r;code = 114 char = o;code = 111 char = n;code = 110 char = o;code = 111 char = l;code = 108 char = o;code = 111 char = g;code = 103 char = i;code = 105 char = e;code = 101 char = s;code = 115 char = ;code = 32 char = o;code = 111 char = f;code = 102 char = ;code = 32 char = a;code = 97 char = n;code = 110 char = c;code = 99 char = i;code = 105 char = e;code = 101 char = n;code = 110 char = t;code = 116 char = ;code = 32 char = r;code = 114 char = i;code = 105 char = b;code = 98 char = o;code = 111 char = s;code = 115 char = o;code = 111 char = m;code = 109 char = a;code = 97 char = l;code = 108 char = ;code = 32 char = e;code = 101 char = v;code = 118 char = o;code = 111 char = l;code = 108 char = u;code = 117 char = t;code = 116 char = i;code = 105 char = o;code = 111 char = n;code = 110 char = .;code = 46 char = ;code = 32 char = T;code = 84 char = h;code = 104 char = e;code = 101 char = ;code = 32 char = m;code = 109 char = e;code = 101 char = t;code = 116 char = h;code = 104 char = o;code = 111 char = d;code = 100 char = ;code = 32 char = u;code = 117 char = s;code = 115 char = e;code = 101 char = s;code = 115 char = ;code = 32 char = s;code = 115 char = t;code = 116 char = r;code = 114 char = u;code = 117 char = c;code = 99 char = t;code = 116 char = u;code = 117 char = r;code = 114 char = e;code = 101 char = -;code = 45 char = b;code = 98 char = a;code = 97 char = s;code = 115 char = e;code = 101 char = d;code = 100 char = ;code = 32 char = a;code = 97 char = n;code = 110 char = d;code = 100 char = ;code = 32 char = s;code = 115 char = e;code = 101 char = q;code = 113 char = u;code = 117 char = e;code = 101 char = n;code = 110 char = c;code = 99 char = e;code = 101 char = -;code = 45 char = b;code = 98 char = a;code = 97 char = s;code = 115 char = e;code = 101 char = d;code = 100 char = ;code = 32 char = c;code = 99 char = o;code = 111 char = m;code = 109 char = p;code = 112 char = a;code = 97 char = r;code = 114 char = i;code = 105 char = s;code = 115 char = o;code = 111 char = n;code = 110 char = ;code = 32 char = o;code = 111 char = f;code = 102 char = ;code = 32 char = t;code = 116 char = h;code = 104 char = e;code = 101 char = ;code = 32 char = l;code = 108 char = a;code = 97 char = r;code = 114 char = g;code = 103 char = e;code = 101 char = ;code = 32 char = s;code = 115 char = u;code = 117 char = b;code = 98 char = u;code = 117 char = n;code = 110 char = i;code = 105 char = t;code = 116 char = s;code = 115 char = ;code = 32 char = (;code = 40 char = L;code = 76 char = S;code = 83 char = U;code = 85 char = s;code = 115 char = );code = 41 char = ;code = 32 char = o;code = 111 char = f;code = 102 char = ;code = 32 char = H;code = 72 char = a;code = 97 char = l;code = 108 char = o;code = 111 char = a;code = 97 char = r;code = 114 char = c;code = 99 char = u;code = 117 char = l;code = 108 char = a;code = 97 char = ;code = 32 char = m;code = 109 char = a;code = 97 char = r;code = 114 char = i;code = 105 char = s;code = 115 char = m;code = 109 char = o;code = 111 char = r;code = 114 char = t;code = 116 char = u;code = 117 char = i;code = 105 char = ;code = 32 char = a;code = 97 char = n;code = 110 char = d;code = 100 char = ;code = 32 char = T;code = 84 char = h;code = 104 char = e;code = 101 char = r;code = 114 char = m;code = 109 char = u;code = 117 char = s;code = 115 char = ;code = 32 char = t;code = 116 char = h;code = 104 char = e;code = 101 char = r;code = 114 char = m;code = 109 char = o;code = 111 char = p;code = 112 char = h;code = 104 char = i;code = 105 char = l;code = 108 char = u;code = 117 char = s;code = 115 char = .;code = 46 char = ;code = 32 char = T;code = 84 char = h;code = 104 char = e;code = 101 char = s;code = 115 char = e;code = 101 char = ;code = 32 char = a;code = 97 char = r;code = 114 char = e;code = 101 char = ;code = 32 char = t;code = 116 char = h;code = 104 char = e;code = 101 char = ;code = 32 char = h;code = 104 char = i;code = 105 char = g;code = 103 char = h;code = 104 char = e;code = 101 char = s;code = 115 char = t;code = 116 char = ;code = 32 char = r;code = 114 char = e;code = 101 char = s;code = 115 char = o;code = 111 char = l;code = 108 char = u;code = 117 char = t;code = 116 char = i;code = 105 char = o;code = 111 char = n;code = 110 char = ;code = 32 char = r;code = 114 char = i;code = 105 char = b;code = 98 char = o;code = 111 char = s;code = 115 char = o;code = 111 char = m;code = 109 char = e;code = 101 char = ;code = 32 char = s;code = 115 char = t;code = 116 char = r;code = 114 char = u;code = 117 char = c;code = 99 char = t;code = 116 char = u;code = 117 char = r;code = 114 char = e;code = 101 char = s;code = 115 char = ;code = 32 char = a;code = 97 char = v;code = 118 char = a;code = 97 char = i;code = 105 char = l;code = 108 char = a;code = 97 char = b;code = 98 char = l;code = 108 char = e;code = 101 char = ;code = 32 char = a;code = 97 char = n;code = 110 char = d;code = 100 char = ;code = 32 char = r;code = 114 char = e;code = 101 char = p;code = 112 char = r;code = 114 char = e;code = 101 char = s;code = 115 char = e;code = 101 char = n;code = 110 char = t;code = 116 char = ;code = 32 char = d;code = 100 char = i;code = 105 char = s;code = 115 char = p;code = 112 char = a;code = 97 char = r;code = 114 char = a;code = 97 char = t;code = 116 char = e;code = 101 char = ;code = 32 char = r;code = 114 char = e;code = 101 char = g;code = 103 char = i;code = 105 char = o;code = 111 char = n;code = 110 char = s;code = 115 char = ;code = 32 char = o;code = 111 char = f;code = 102 char = ;code = 32 char = t;code = 116 char = h;code = 104 char = e;code = 101 char = ;code = 32 char = e;code = 101 char = v;code = 118 char = o;code = 111 char = l;code = 108 char = u;code = 117 char = t;code = 116 char = i;code = 105 char = o;code = 111 char = n;code = 110 char = a;code = 97 char = r;code = 114 char = y;code = 121 char = ;code = 32 char = t;code = 116 char = r;code = 114 char = e;code = 101 char = e;code = 101 char = .;code = 46 char = ;code = 32 char = W;code = 87 char = e;code = 101 char = ;code = 32 char = h;code = 104 char = a;code = 97 char = v;code = 118 char = e;code = 101 char = ;code = 32 char = s;code = 115 char = e;code = 101 char = c;code = 99 char = t;code = 116 char = i;code = 105 char = o;code = 111 char = n;code = 110 char = e;code = 101 char = d;code = 100 char = ;code = 32 char = t;code = 116 char = h;code = 104 char = e;code = 101 char = ;code = 32 char = s;code = 115 char = u;code = 117 char = p;code = 112 char = e;code = 101 char = r;code = 114 char = i;code = 105 char = m;code = 109 char = p;code = 112 char = o;code = 111 char = s;code = 115 char = e;code = 101 char = d;code = 100 char = ;code = 32 char = L;code = 76 char = S;code = 83 char = U;code = 85 char = s;code = 115 char = ;code = 32 char = i;code = 105 char = n;code = 110 char = t;code = 116 char = o;code = 111 char = ;code = 32 char = c;code = 99 char = o;code = 111 char = n;code = 110 char = c;code = 99 char = e;code = 101 char = n;code = 110 char = t;code = 116 char = r;code = 114 char = i;code = 105 char = c;code = 99 char = ;code = 32 char = s;code = 115 char = h;code = 104 char = e;code = 101 char = l;code = 108 char = l;code = 108 char = s;code = 115 char = ,;code = 44 char = ;code = 32 char = l;code = 108 char = i;code = 105 char = k;code = 107 char = e;code = 101 char = ;code = 32 char = a;code = 97 char = n;code = 110 char = ;code = 32 char = o;code = 111 char = n;code = 110 char = i;code = 105 char = o;code = 111 char = n;code = 110 char = ,;code = 44 char = ;code = 32 char = u;code = 117 char = s;code = 115 char = i;code = 105 char = n;code = 110 char = g;code = 103 char = ;code = 32 char = t;code = 116 char = h;code = 104 char = e;code = 101 char = ;code = 32 char = s;code = 115 char = i;code = 105 char = t;code = 116 char = e;code = 101 char = ;code = 32 char = o;code = 111 char = f;code = 102 char = ;code = 32 char = p;code = 112 char = e;code = 101 char = p;code = 112 char = t;code = 116 char = i;code = 105 char = d;code = 100 char = y;code = 121 char = l;code = 108 char = ;code = 32 char = t;code = 116 char = r;code = 114 char = a;code = 97 char = n;code = 110 char = s;code = 115 char = f;code = 102 char = e;code = 101 char = r;code = 114 char = ;code = 32 char = a;code = 97 char = s;code = 115 char = ;code = 32 char = t;code = 116 char = h;code = 104 char = e;code = 101 char = ;code = 32 char = o;code = 111 char = r;code = 114 char = i;code = 105 char = g;code = 103 char = i;code = 105 char = n;code = 110 char = ;code = 32 char = (;code = 40 char = t;code = 116 char = h;code = 104 char = e;code = 101 char = ;code = 32 char = P;code = 80 char = T;code = 84 char = -;code = 45 char = o;code = 111 char = r;code = 114 char = i;code = 105 char = g;code = 103 char = i;code = 105 char = n;code = 110 char = );code = 41 char = .;code = 46 char = ;code = 32 char = T;code = 84 char = h;code = 104 char = i;code = 105 char = s;code = 115 char = ;code = 32 char = s;code = 115 char = p;code = 112 char = h;code = 104 char = e;code = 101 char = r;code = 114 char = i;code = 105 char = c;code = 99 char = a;code = 97 char = l;code = 108 char = ;code = 32 char = a;code = 97 char = p;code = 112 char = p;code = 112 char = r;code = 114 char = o;code = 111 char = x;code = 120 char = i;code = 105 char = m;code = 109 char = a;code = 97 char = t;code = 116 char = i;code = 105 char = o;code = 111 char = n;code = 110 char = ;code = 32 char = c;code = 99 char = o;code = 111 char = m;code = 109 char = b;code = 98 char = i;code = 105 char = n;code = 110 char = e;code = 101 char = d;code = 100 char = ;code = 32 char = w;code = 119 char = i;code = 105 char = t;code = 116 char = h;code = 104 char = ;code = 32 char = a;code = 97 char = ;code = 32 char = s;code = 115 char = h;code = 104 char = e;code = 101 char = l;code = 108 char = l;code = 108 char = -;code = 45 char = b;code = 98 char = y;code = 121 char = -;code = 45 char = s;code = 115 char = h;code = 104 char = e;code = 101 char = l;code = 108 char = l;code = 108 char = ;code = 32 char = c;code = 99 char = o;code = 111 char = m;code = 109 char = p;code = 112 char = a;code = 97 char = r;code = 114 char = i;code = 105 char = s;code = 115 char = o;code = 111 char = n;code = 110 char = ;code = 32 char = c;code = 99 char = a;code = 97 char = p;code = 112 char = t;code = 116 char = u;code = 117 char = r;code = 114 char = e;code = 101 char = s;code = 115 char = ;code = 32 char = s;code = 115 char = i;code = 105 char = g;code = 103 char = n;code = 110 char = i;code = 105 char = f;code = 102 char = i;code = 105 char = c;code = 99 char = a;code = 97 char = n;code = 110 char = t;code = 116 char = ;code = 32 char = i;code = 105 char = n;code = 110 char = f;code = 102 char = o;code = 111 char = r;code = 114 char = m;code = 109 char = a;code = 97 char = t;code = 116 char = i;code = 105 char = o;code = 111 char = n;code = 110 char = ;code = 32 char = a;code = 97 char = l;code = 108 char = o;code = 111 char = n;code = 110 char = g;code = 103 char = ;code = 32 char = t;code = 116 char = h;code = 104 char = e;code = 101 char = ;code = 32 char = e;code = 101 char = v;code = 118 char = o;code = 111 char = l;code = 108 char = u;code = 117 char = t;code = 116 char = i;code = 105 char = o;code = 111 char = n;code = 110 char = a;code = 97 char = r;code = 114 char = y;code = 121 char = ;code = 32 char = t;code = 116 char = i;code = 105 char = m;code = 109 char = e;code = 101 char = ;code = 32 char = l;code = 108 char = i;code = 105 char = n;code = 110 char = e;code = 101 char = ;code = 32 char = r;code = 114 char = e;code = 101 char = v;code = 118 char = e;code = 101 char = a;code = 97 char = l;code = 108 char = i;code = 105 char = n;code = 110 char = g;code = 103 char = ,;code = 44 char = ;code = 32 char = f;code = 102 char = o;code = 111 char = r;code = 114 char = ;code = 32 char = e;code = 101 char = x;code = 120 char = a;code = 97 char = m;code = 109 char = p;code = 112 char = l;code = 108 char = e;code = 101 char = ,;code = 44 char = ;code = 32 char = t;code = 116 char = h;code = 104 char = a;code = 97 char = t;code = 116 char = ;code = 32 char = s;code = 115 char = e;code = 101 char = q;code = 113 char = u;code = 117 char = e;code = 101 char = n;code = 110 char = c;code = 99 char = e;code = 101 char = ;code = 32 char = a;code = 97 char = n;code = 110 char = d;code = 100 char = ;code = 32 char = c;code = 99 char = o;code = 111 char = n;code = 110 char = f;code = 102 char = o;code = 111 char = r;code = 114 char = m;code = 109 char = a;code = 97 char = t;code = 116 char = i;code = 105 char = o;code = 111 char = n;code = 110 char = a;code = 97 char = l;code = 108 char = ;code = 32 char = s;code = 115 char = i;code = 105 char = m;code = 109 char = i;code = 105 char = l;code = 108 char = a;code = 97 char = r;code = 114 char = i;code = 105 char = t;code = 116 char = y;code = 121 char = ;code = 32 char = o;code = 111 char = f;code = 102 char = ;code = 32 char = t;code = 116 char = h;code = 104 char = e;code = 101 char = ;code = 32 char = 2;code = 50 char = 3;code = 51 char = S;code = 83 char = ;code = 32 char = r;code = 114 char = R;code = 82 char = N;code = 78 char = A;code = 65 char = s;code = 115 char = ;code = 32 char = a;code = 97 char = r;code = 114 char = e;code = 101 char = ;code = 32 char = g;code = 103 char = r;code = 114 char = e;code = 101 char = a;code = 97 char = t;code = 116 char = e;code = 101 char = s;code = 115 char = t;code = 116 char = ;code = 32 char = n;code = 110 char = e;code = 101 char = a;code = 97 char = r;code = 114 char = ;code = 32 char = t;code = 116 char = h;code = 104 char = e;code = 101 char = ;code = 32 char = P;code = 80 char = T;code = 84 char = -;code = 45 char = o;code = 111 char = r;code = 114 char = i;code = 105 char = g;code = 103 char = i;code = 105 char = n;code = 110 char = ;code = 32 char = a;code = 97 char = n;code = 110 char = d;code = 100 char = ;code = 32 char = d;code = 100 char = i;code = 105 char = v;code = 118 char = e;code = 101 char = r;code = 114 char = g;code = 103 char = e;code = 101 char = ;code = 32 char = s;code = 115 char = m;code = 109 char = o;code = 111 char = o;code = 111 char = t;code = 116 char = h;code = 104 char = l;code = 108 char = y;code = 121 char = ;code = 32 char = w;code = 119 char = i;code = 105 char = t;code = 116 char = h;code = 104 char = ;code = 32 char = d;code = 100 char = i;code = 105 char = s;code = 115 char = t;code = 116 char = a;code = 97 char = n;code = 110 char = c;code = 99 char = e;code = 101 char = ;code = 32 char = f;code = 102 char = r;code = 114 char = o;code = 111 char = m;code = 109 char = ;code = 32 char = i;code = 105 char = t;code = 116 char = .;code = 46 char = ;code = 32 char = T;code = 84 char = h;code = 104 char = e;code = 101 char = ;code = 32 char = r;code = 114 char = e;code = 101 char = s;code = 115 char = u;code = 117 char = l;code = 108 char = t;code = 116 char = s;code = 115 char = ;code = 32 char = s;code = 115 char = u;code = 117 char = g;code = 103 char = g;code = 103 char = e;code = 101 char = s;code = 115 char = t;code = 116 char = ;code = 32 char = t;code = 116 char = h;code = 104 char = a;code = 97 char = t;code = 116 char = ;code = 32 char = t;code = 116 char = h;code = 104 char = e;code = 101 char = ;code = 32 char = c;code = 99 char = o;code = 111 char = n;code = 110 char = f;code = 102 char = o;code = 111 char = r;code = 114 char = m;code = 109 char = a;code = 97 char = t;code = 116 char = i;code = 105 char = o;code = 111 char = n;code = 110 char = ;code = 32 char = a;code = 97 char = n;code = 110 char = d;code = 100 char = ;code = 32 char = i;code = 105 char = n;code = 110 char = t;code = 116 char = e;code = 101 char = r;code = 114 char = a;code = 97 char = c;code = 99 char = t;code = 116 char = i;code = 105 char = o;code = 111 char = n;code = 110 char = s;code = 115 char = ;code = 32 char = o;code = 111 char = f;code = 102 char = ;code = 32 char = b;code = 98 char = o;code = 111 char = t;code = 116 char = h;code = 104 char = ;code = 32 char = R;code = 82 char = N;code = 78 char = A;code = 65 char = ;code = 32 char = a;code = 97 char = n;code = 110 char = d;code = 100 char = ;code = 32 char = p;code = 112 char = r;code = 114 char = o;code = 111 char = t;code = 116 char = e;code = 101 char = i;code = 105 char = n;code = 110 char = ;code = 32 char = c;code = 99 char = a;code = 97 char = n;code = 110 char = ;code = 32 char = b;code = 98 char = e;code = 101 char = ;code = 32 char = d;code = 100 char = e;code = 101 char = s;code = 115 char = c;code = 99 char = r;code = 114 char = i;code = 105 char = b;code = 98 char = e;code = 101 char = d;code = 100 char = ;code = 32 char = a;code = 97 char = s;code = 115 char = ;code = 32 char = c;code = 99 char = h;code = 104 char = a;code = 97 char = n;code = 110 char = g;code = 103 char = i;code = 105 char = n;code = 110 char = g;code = 103 char = ,;code = 44 char = ;code = 32 char = i;code = 105 char = n;code = 110 char = ;code = 32 char = a;code = 97 char = n;code = 110 char = ;code = 32 char = o;code = 111 char = b;code = 98 char = s;code = 115 char = e;code = 101 char = r;code = 114 char = v;code = 118 char = a;code = 97 char = b;code = 98 char = l;code = 108 char = e;code = 101 char = ;code = 32 char = m;code = 109 char = a;code = 97 char = n;code = 110 char = n;code = 110 char = e;code = 101 char = r;code = 114 char = ,;code = 44 char = ;code = 32 char = o;code = 111 char = v;code = 118 char = e;code = 101 char = r;code = 114 char = ;code = 32 char = e;code = 101 char = v;code = 118 char = o;code = 111 char = l;code = 108 char = u;code = 117 char = t;code = 116 char = i;code = 105 char = o;code = 111 char = n;code = 110 char = a;code = 97 char = r;code = 114 char = y;code = 121 char = ;code = 32 char = t;code = 116 char = i;code = 105 char = m;code = 109 char = e;code = 101 char = .;code = 46 char = ;code = 32 char = T;code = 84 char = h;code = 104 char = e;code = 101 char = ;code = 32 char = t;code = 116 char = e;code = 101 char = n;code = 110 char = d;code = 100 char = e;code = 101 char = n;code = 110 char = c;code = 99 char = y;code = 121 char = ;code = 32 char = o;code = 111 char = f;code = 102 char = ;code = 32 char = m;code = 109 char = a;code = 97 char = c;code = 99 char = r;code = 114 char = o;code = 111 char = m;code = 109 char = o;code = 111 char = l;code = 108 char = e;code = 101 char = c;code = 99 char = u;code = 117 char = l;code = 108 char = e;code = 101 char = s;code = 115 char = ;code = 32 char = t;code = 116 char = o;code = 111 char = ;code = 32 char = a;code = 97 char = s;code = 115 char = s;code = 115 char = u;code = 117 char = m;code = 109 char = e;code = 101 char = ;code = 32 char = r;code = 114 char = e;code = 101 char = g;code = 103 char = u;code = 117 char = l;code = 108 char = a;code = 97 char = r;code = 114 char = ;code = 32 char = s;code = 115 char = e;code = 101 char = c;code = 99 char = o;code = 111 char = n;code = 110 char = d;code = 100 char = a;code = 97 char = r;code = 114 char = y;code = 121 char = ;code = 32 char = s;code = 115 char = t;code = 116 char = r;code = 114 char = u;code = 117 char = c;code = 99 char = t;code = 116 char = u;code = 117 char = r;code = 114 char = a;code = 97 char = l;code = 108 char = ;code = 32 char = e;code = 101 char = l;code = 108 char = e;code = 101 char = m;code = 109 char = e;code = 101 char = n;code = 110 char = t;code = 116 char = s;code = 115 char = ;code = 32 char = s;code = 115 char = u;code = 117 char = c;code = 99 char = h;code = 104 char = ;code = 32 char = a;code = 97 char = s;code = 115 char = ;code = 32 char = A;code = 65 char = -;code = 45 char = f;code = 102 char = o;code = 111 char = r;code = 114 char = m;code = 109 char = ;code = 32 char = h;code = 104 char = e;code = 101 char = l;code = 108 char = i;code = 105 char = c;code = 99 char = e;code = 101 char = s;code = 115 char = ;code = 32 char = w;code = 119 char = i;code = 105 char = t;code = 116 char = h;code = 104 char = ;code = 32 char = W;code = 87 char = a;code = 97 char = t;code = 116 char = s;code = 115 char = o;code = 111 char = n;code = 110 char = -;code = 45 char = C;code = 67 char = r;code = 114 char = i;code = 105 char = c;code = 99 char = k;code = 107 char = ;code = 32 char = b;code = 98 char = a;code = 97 char = s;code = 115 char = e;code = 101 char = ;code = 32 char = p;code = 112 char = a;code = 97 char = i;code = 105 char = r;code = 114 char = s;code = 115 char = ;code = 32 char = (;code = 40 char = R;code = 82 char = N;code = 78 char = A;code = 65 char = );code = 41 char = ;code = 32 char = a;code = 97 char = n;code = 110 char = d;code = 100 char = ;code = 32 char = a;code = 97 char = l;code = 108 char = p;code = 112 char = h;code = 104 char = a;code = 97 char = -;code = 45 char = h;code = 104 char = e;code = 101 char = l;code = 108 char = i;code = 105 char = c;code = 99 char = e;code = 101 char = s;code = 115 char = ;code = 32 char = a;code = 97 char = n;code = 110 char = d;code = 100 char = ;code = 32 char = b;code = 98 char = e;code = 101 char = t;code = 116 char = a;code = 97 char = -;code = 45 char = s;code = 115 char = h;code = 104 char = e;code = 101 char = e;code = 101 char = t;code = 116 char = s;code = 115 char = ;code = 32 char = (;code = 40 char = p;code = 112 char = r;code = 114 char = o;code = 111 char = t;code = 116 char = e;code = 101 char = i;code = 105 char = n;code = 110 char = );code = 41 char = ;code = 32 char = i;code = 105 char = s;code = 115 char = ;code = 32 char = l;code = 108 char = o;code = 111 char = w;code = 119 char = ;code = 32 char = a;code = 97 char = t;code = 116 char = ;code = 32 char = e;code = 101 char = a;code = 97 char = r;code = 114 char = l;code = 108 char = y;code = 121 char = ;code = 32 char = t;code = 116 char = i;code = 105 char = m;code = 109 char = e;code = 101 char = ;code = 32 char = p;code = 112 char = o;code = 111 char = i;code = 105 char = n;code = 110 char = t;code = 116 char = s;code = 115 char = ;code = 32 char = b;code = 98 char = u;code = 117 char = t;code = 116 char = ;code = 32 char = i;code = 105 char = n;code = 110 char = c;code = 99 char = r;code = 114 char = e;code = 101 char = a;code = 97 char = s;code = 115 char = e;code = 101 char = s;code = 115 char = ;code = 32 char = a;code = 97 char = s;code = 115 char = ;code = 32 char = t;code = 116 char = i;code = 105 char = m;code = 109 char = e;code = 101 char = ;code = 32 char = p;code = 112 char = r;code = 114 char = o;code = 111 char = g;code = 103 char = r;code = 114 char = e;code = 101 char = s;code = 115 char = s;code = 115 char = e;code = 101 char = s;code = 115 char = .;code = 46 char = ;code = 32 char = T;code = 84 char = h;code = 104 char = e;code = 101 char = ;code = 32 char = c;code = 99 char = o;code = 111 char = n;code = 110 char = f;code = 102 char = o;code = 111 char = r;code = 114 char = m;code = 109 char = a;code = 97 char = t;code = 116 char = i;code = 105 char = o;code = 111 char = n;code = 110 char = s;code = 115 char = ;code = 32 char = o;code = 111 char = f;code = 102 char = ;code = 32 char = r;code = 114 char = i;code = 105 char = b;code = 98 char = o;code = 111 char = s;code = 115 char = o;code = 111 char = m;code = 109 char = a;code = 97 char = l;code = 108 char = ;code = 32 char = p;code = 112 char = r;code = 114 char = o;code = 111 char = t;code = 116 char = e;code = 101 char = i;code = 105 char = n;code = 110 char = ;code = 32 char = c;code = 99 char = o;code = 111 char = m;code = 109 char = p;code = 112 char = o;code = 111 char = n;code = 110 char = e;code = 101 char = n;code = 110 char = t;code = 116 char = s;code = 115 char = ;code = 32 char = n;code = 110 char = e;code = 101 char = a;code = 97 char = r;code = 114 char = ;code = 32 char = t;code = 116 char = h;code = 104 char = e;code = 101 char = ;code = 32 char = P;code = 80 char = T;code = 84 char = -;code = 45 char = o;code = 111 char = r;code = 114 char = i;code = 105 char = g;code = 103 char = i;code = 105 char = n;code = 110 char = ;code = 32 char = s;code = 115 char = u;code = 117 char = g;code = 103 char = g;code = 103 char = e;code = 101 char = s;code = 115 char = t;code = 116 char = ;code = 32 char = t;code = 116 char = h;code = 104 char = a;code = 97 char = t;code = 116 char = ;code = 32 char = t;code = 116 char = h;code = 104 char = e;code = 101 char = y;code = 121 char = ;code = 32 char = m;code = 109 char = a;code = 97 char = y;code = 121 char = ;code = 32 char = b;code = 98 char = e;code = 101 char = ;code = 32 char = m;code = 109 char = o;code = 111 char = l;code = 108 char = e;code = 101 char = c;code = 99 char = u;code = 117 char = l;code = 108 char = a;code = 97 char = r;code = 114 char = ;code = 32 char = f;code = 102 char = o;code = 111 char = s;code = 115 char = s;code = 115 char = i;code = 105 char = l;code = 108 char = s;code = 115 char = ;code = 32 char = o;code = 111 char = f;code = 102 char = ;code = 32 char = t;code = 116 char = h;code = 104 char = e;code = 101 char = ;code = 32 char = p;code = 112 char = e;code = 101 char = p;code = 112 char = t;code = 116 char = i;code = 105 char = d;code = 100 char = e;code = 101 char = ;code = 32 char = a;code = 97 char = n;code = 110 char = c;code = 99 char = e;code = 101 char = s;code = 115 char = t;code = 116 char = o;code = 111 char = r;code = 114 char = s;code = 115 char = ;code = 32 char = o;code = 111 char = f;code = 102 char = ;code = 32 char = r;code = 114 char = i;code = 105 char = b;code = 98 char = o;code = 111 char = s;code = 115 char = o;code = 111 char = m;code = 109 char = a;code = 97 char = l;code = 108 char = ;code = 32 char = p;code = 112 char = r;code = 114 char = o;code = 111 char = t;code = 116 char = e;code = 101 char = i;code = 105 char = n;code = 110 char = s;code = 115 char = .;code = 46 char = ;code = 32 char = T;code = 84 char = h;code = 104 char = e;code = 101 char = i;code = 105 char = r;code = 114 char = ;code = 32 char = a;code = 97 char = b;code = 98 char = b;code = 98 char = r;code = 114 char = e;code = 101 char = v;code = 118 char = i;code = 105 char = a;code = 97 char = t;code = 116 char = e;code = 101 char = d;code = 100 char = ;code = 32 char = l;code = 108 char = e;code = 101 char = n;code = 110 char = g;code = 103 char = t;code = 116 char = h;code = 104 char = ;code = 32 char = m;code = 109 char = a;code = 97 char = y;code = 121 char = ;code = 32 char = h;code = 104 char = a;code = 97 char = v;code = 118 char = e;code = 101 char = ;code = 32 char = p;code = 112 char = r;code = 114 char = o;code = 111 char = s;code = 115 char = c;code = 99 char = r;code = 114 char = i;code = 105 char = b;code = 98 char = e;code = 101 char = d;code = 100 char = ;code = 32 char = f;code = 102 char = o;code = 111 char = r;code = 114 char = m;code = 109 char = a;code = 97 char = t;code = 116 char = i;code = 105 char = o;code = 111 char = n;code = 110 char = ;code = 32 char = o;code = 111 char = f;code = 102 char = ;code = 32 char = s;code = 115 char = e;code = 101 char = c;code = 99 char = o;code = 111 char = n;code = 110 char = d;code = 100 char = a;code = 97 char = r;code = 114 char = y;code = 121 char = ;code = 32 char = s;code = 115 char = t;code = 116 char = r;code = 114 char = u;code = 117 char = c;code = 99 char = t;code = 116 char = u;code = 117 char = r;code = 114 char = e;code = 101 char = ,;code = 44 char = ;code = 32 char = w;code = 119 char = h;code = 104 char = i;code = 105 char = c;code = 99 char = h;code = 104 char = ;code = 32 char = i;code = 105 char = s;code = 115 char = ;code = 32 char = i;code = 105 char = n;code = 110 char = d;code = 100 char = e;code = 101 char = e;code = 101 char = d;code = 100 char = ;code = 32 char = n;code = 110 char = e;code = 101 char = a;code = 97 char = r;code = 114 char = l;code = 108 char = y;code = 121 char = ;code = 32 char = a;code = 97 char = b;code = 98 char = s;code = 115 char = e;code = 101 char = n;code = 110 char = t;code = 116 char = ;code = 32 char = f;code = 102 char = r;code = 114 char = o;code = 111 char = m;code = 109 char = ;code = 32 char = t;code = 116 char = h;code = 104 char = e;code = 101 char = ;code = 32 char = r;code = 114 char = e;code = 101 char = g;code = 103 char = i;code = 105 char = o;code = 111 char = n;code = 110 char = ;code = 32 char = o;code = 111 char = f;code = 102 char = ;code = 32 char = t;code = 116 char = h;code = 104 char = e;code = 101 char = ;code = 32 char = L;code = 76 char = S;code = 83 char = U;code = 85 char = ;code = 32 char = n;code = 110 char = e;code = 101 char = a;code = 97 char = r;code = 114 char = e;code = 101 char = s;code = 115 char = t;code = 116 char = ;code = 32 char = t;code = 116 char = h;code = 104 char = e;code = 101 char = ;code = 32 char = P;code = 80 char = T;code = 84 char = -;code = 45 char = o;code = 111 char = r;code = 114 char = i;code = 105 char = g;code = 103 char = i;code = 105 char = n;code = 110 char = .;code = 46 char = ;code = 32 char = F;code = 70 char = o;code = 111 char = r;code = 114 char = m;code = 109 char = a;code = 97 char = t;code = 116 char = i;code = 105 char = o;code = 111 char = n;code = 110 char = ;code = 32 char = a;code = 97 char = n;code = 110 char = d;code = 100 char = ;code = 32 char = e;code = 101 char = v;code = 118 char = o;code = 111 char = l;code = 108 char = u;code = 117 char = t;code = 116 char = i;code = 105 char = o;code = 111 char = n;code = 110 char = ;code = 32 char = o;code = 111 char = f;code = 102 char = ;code = 32 char = t;code = 116 char = h;code = 104 char = e;code = 101 char = ;code = 32 char = e;code = 101 char = a;code = 97 char = r;code = 114 char = l;code = 108 char = y;code = 121 char = ;code = 32 char = P;code = 80 char = T;code = 84 char = ;code = 32 char = c;code = 99 char = e;code = 101 char = n;code = 110 char = t;code = 116 char = e;code = 101 char = r;code = 114 char = ;code = 32 char = m;code = 109 char = a;code = 97 char = y;code = 121 char = ;code = 32 char = h;code = 104 char = a;code = 97 char = v;code = 118 char = e;code = 101 char = ;code = 32 char = i;code = 105 char = n;code = 110 char = v;code = 118 char = o;code = 111 char = l;code = 108 char = v;code = 118 char = e;code = 101 char = d;code = 100 char = ;code = 32 char = M;code = 77 char = g;code = 103 char = (;code = 40 char = 2;code = 50 char = +;code = 43 char = );code = 41 char = -;code = 45 char = m;code = 109 char = e;code = 101 char = d;code = 100 char = i;code = 105 char = a;code = 97 char = t;code = 116 char = e;code = 101 char = d;code = 100 char = ;code = 32 char = a;code = 97 char = s;code = 115 char = s;code = 115 char = e;code = 101 char = m;code = 109 char = b;code = 98 char = l;code = 108 char = y;code = 121 char = ;code = 32 char = o;code = 111 char = f;code = 102 char = ;code = 32 char = a;code = 97 char = t;code = 116 char = ;code = 32 char = l;code = 108 char = e;code = 101 char = a;code = 97 char = s;code = 115 char = t;code = 116 char = ;code = 32 char = p;code = 112 char = a;code = 97 char = r;code = 114 char = t;code = 116 char = i;code = 105 char = a;code = 97 char = l;code = 108 char = l;code = 108 char = y;code = 121 char = ;code = 32 char = s;code = 115 char = i;code = 105 char = n;code = 110 char = g;code = 103 char = l;code = 108 char = e;code = 101 char = -;code = 45 char = s;code = 115 char = t;code = 116 char = r;code = 114 char = a;code = 97 char = n;code = 110 char = d;code = 100 char = e;code = 101 char = d;code = 100 char = ;code = 32 char = R;code = 82 char = N;code = 78 char = A;code = 65 char = ;code = 32 char = o;code = 111 char = l;code = 108 char = i;code = 105 char = g;code = 103 char = o;code = 111 char = m;code = 109 char = e;code = 101 char = r;code = 114 char = s;code = 115 char = ;code = 32 char = o;code = 111 char = r;code = 114 char = ;code = 32 char = p;code = 112 char = o;code = 111 char = l;code = 108 char = y;code = 121 char = m;code = 109 char = e;code = 101 char = r;code = 114 char = s;code = 115 char = .;code = 46 char = ;code = 32 char = A;code = 65 char = s;code = 115 char = ;code = 32 char = o;code = 111 char = n;code = 110 char = e;code = 101 char = ;code = 32 char = m;code = 109 char = o;code = 111 char = v;code = 118 char = e;code = 101 char = s;code = 115 char = ;code = 32 char = f;code = 102 char = r;code = 114 char = o;code = 111 char = m;code = 109 char = ;code = 32 char = c;code = 99 char = e;code = 101 char = n;code = 110 char = t;code = 116 char = e;code = 101 char = r;code = 114 char = ;code = 32 char = t;code = 116 char = o;code = 111 char = ;code = 32 char = p;code = 112 char = e;code = 101 char = r;code = 114 char = i;code = 105 char = p;code = 112 char = h;code = 104 char = e;code = 101 char = r;code = 114 char = y;code = 121 char = ,;code = 44 char = ;code = 32 char = p;code = 112 char = r;code = 114 char = o;code = 111 char = t;code = 116 char = e;code = 101 char = i;code = 105 char = n;code = 110 char = s;code = 115 char = ;code = 32 char = a;code = 97 char = p;code = 112 char = p;code = 112 char = e;code = 101 char = a;code = 97 char = r;code = 114 char = ;code = 32 char = t;code = 116 char = o;code = 111 char = ;code = 32 char = r;code = 114 char = e;code = 101 char = p;code = 112 char = l;code = 108 char = a;code = 97 char = c;code = 99 char = e;code = 101 char = ;code = 32 char = m;code = 109 char = a;code = 97 char = g;code = 103 char = n;code = 110 char = e;code = 101 char = s;code = 115 char = i;code = 105 char = u;code = 117 char = m;code = 109 char = ;code = 32 char = i;code = 105 char = o;code = 111 char = n;code = 110 char = s;code = 115 char = .;code = 46 char = ;code = 32 char = T;code = 84 char = h;code = 104 char = e;code = 101 char = ;code = 32 char = L;code = 76 char = S;code = 83 char = U;code = 85 char = ;code = 32 char = i;code = 105 char = s;code = 115 char = ;code = 32 char = k;code = 107 char = n;code = 110 char = o;code = 111 char = w;code = 119 char = n;code = 110 char = ;code = 32 char = t;code = 116 char = o;code = 111 char = ;code = 32 char = h;code = 104 char = a;code = 97 char = v;code = 118 char = e;code = 101 char = ;code = 32 char = u;code = 117 char = n;code = 110 char = d;code = 100 char = e;code = 101 char = r;code = 114 char = g;code = 103 char = o;code = 111 char = n;code = 110 char = e;code = 101 char = ;code = 32 char = l;code = 108 char = a;code = 97 char = r;code = 114 char = g;code = 103 char = e;code = 101 char = -;code = 45 char = s;code = 115 char = c;code = 99 char = a;code = 97 char = l;code = 108 char = e;code = 101 char = ;code = 32 char = c;code = 99 char = o;code = 111 char = n;code = 110 char = f;code = 102 char = o;code = 111 char = r;code = 114 char = m;code = 109 char = a;code = 97 char = t;code = 116 char = i;code = 105 char = o;code = 111 char = n;code = 110 char = ;code = 32 char = c;code = 99 char = h;code = 104 char = a;code = 97 char = n;code = 110 char = g;code = 103 char = e;code = 101 char = s;code = 115 char = ;code = 32 char = u;code = 117 char = p;code = 112 char = o;code = 111 char = n;code = 110 char = ;code = 32 char = a;code = 97 char = s;code = 115 char = s;code = 115 char = e;code = 101 char = m;code = 109 char = b;code = 98 char = l;code = 108 char = y;code = 121 char = .;code = 46 char = ;code = 32 char = T;code = 84 char = h;code = 104 char = e;code = 101 char = ;code = 32 char = T;code = 84 char = .;code = 46 char = ;code = 32 char = t;code = 116 char = h;code = 104 char = e;code = 101 char = r;code = 114 char = m;code = 109 char = o;code = 111 char = p;code = 112 char = h;code = 104 char = i;code = 105 char = l;code = 108 char = u;code = 117 char = s;code = 115 char = ;code = 32 char = L;code = 76 char = S;code = 83 char = U;code = 85 char = ;code = 32 char = a;code = 97 char = n;code = 110 char = a;code = 97 char = l;code = 108 char = y;code = 121 char = z;code = 122 char = e;code = 101 char = d;code = 100 char = ;code = 32 char = h;code = 104 char = e;code = 101 char = r;code = 114 char = e;code = 101 char = ;code = 32 char = i;code = 105 char = s;code = 115 char = ;code = 32 char = p;code = 112 char = a;code = 97 char = r;code = 114 char = t;code = 116 char = ;code = 32 char = o;code = 111 char = f;code = 102 char = ;code = 32 char = a;code = 97 char = ;code = 32 char = f;code = 102 char = u;code = 117 char = l;code = 108 char = l;code = 108 char = y;code = 121 char = ;code = 32 char = a;code = 97 char = s;code = 115 char = s;code = 115 char = e;code = 101 char = m;code = 109 char = b;code = 98 char = l;code = 108 char = e;code = 101 char = d;code = 100 char = ;code = 32 char = r;code = 114 char = i;code = 105 char = b;code = 98 char = o;code = 111 char = s;code = 115 char = o;code = 111 char = m;code = 109 char = e;code = 101 char = ,;code = 44 char = ;code = 32 char = w;code = 119 char = h;code = 104 char = e;code = 101 char = r;code = 114 char = e;code = 101 char = a;code = 97 char = s;code = 115 char = ;code = 32 char = t;code = 116 char = h;code = 104 char = e;code = 101 char = ;code = 32 char = H;code = 72 char = .;code = 46 char = ;code = 32 char = m;code = 109 char = a;code = 97 char = r;code = 114 char = i;code = 105 char = s;code = 115 char = m;code = 109 char = o;code = 111 char = r;code = 114 char = t;code = 116 char = u;code = 117 char = i;code = 105 char = ;code = 32 char = L;code = 76 char = S;code = 83 char = U;code = 85 char = ;code = 32 char = a;code = 97 char = n;code = 110 char = a;code = 97 char = l;code = 108 char = y;code = 121 char = z;code = 122 char = e;code = 101 char = d;code = 100 char = ;code = 32 char = h;code = 104 char = e;code = 101 char = r;code = 114 char = e;code = 101 char = ;code = 32 char = i;code = 105 char = s;code = 115 char = ;code = 32 char = d;code = 100 char = i;code = 105 char = s;code = 115 char = s;code = 115 char = o;code = 111 char = c;code = 99 char = i;code = 105 char = a;code = 97 char = t;code = 116 char = e;code = 101 char = d;code = 100 char = ;code = 32 char = f;code = 102 char = r;code = 114 char = o;code = 111 char = m;code = 109 char = ;code = 32 char = o;code = 111 char = t;code = 116 char = h;code = 104 char = e;code = 101 char = r;code = 114 char = ;code = 32 char = r;code = 114 char = i;code = 105 char = b;code = 98 char = o;code = 111 char = s;code = 115 char = o;code = 111 char = m;code = 109 char = a;code = 97 char = l;code = 108 char = ;code = 32 char = c;code = 99 char = o;code = 111 char = m;code = 109 char = p;code = 112 char = o;code = 111 char = n;code = 110 char = e;code = 101 char = n;code = 110 char = t;code = 116 char = s;code = 115 char = .;code = 46 char = ;code = 32 char = L;code = 76 char = a;code = 97 char = r;code = 114 char = g;code = 103 char = e;code = 101 char = -;code = 45 char = s;code = 115 char = c;code = 99 char = a;code = 97 char = l;code = 108 char = e;code = 101 char = ;code = 32 char = c;code = 99 char = o;code = 111 char = n;code = 110 char = f;code = 102 char = o;code = 111 char = r;code = 114 char = m;code = 109 char = a;code = 97 char = t;code = 116 char = i;code = 105 char = o;code = 111 char = n;code = 110 char = a;code = 97 char = l;code = 108 char = ;code = 32 char = d;code = 100 char = i;code = 105 char = f;code = 102 char = f;code = 102 char = e;code = 101 char = r;code = 114 char = e;code = 101 char = n;code = 110 char = c;code = 99 char = e;code = 101 char = s;code = 115 char = ;code = 32 char = i;code = 105 char = n;code = 110 char = ;code = 32 char = t;code = 116 char = h;code = 104 char = e;code = 101 char = ;code = 32 char = 2;code = 50 char = 3;code = 51 char = S;code = 83 char = ;code = 32 char = r;code = 114 char = R;code = 82 char = N;code = 78 char = A;code = 65 char = s;code = 115 char = ;code = 32 char = a;code = 97 char = r;code = 114 char = e;code = 101 char = ;code = 32 char = e;code = 101 char = v;code = 118 char = i;code = 105 char = d;code = 100 char = e;code = 101 char = n;code = 110 char = t;code = 116 char = ;code = 32 char = f;code = 102 char = r;code = 114 char = o;code = 111 char = m;code = 109 char = ;code = 32 char = s;code = 115 char = u;code = 117 char = p;code = 112 char = e;code = 101 char = r;code = 114 char = i;code = 105 char = m;code = 109 char = p;code = 112 char = o;code = 111 char = s;code = 115 char = i;code = 105 char = t;code = 116 char = i;code = 105 char = o;code = 111 char = n;code = 110 char = ;code = 32 char = a;code = 97 char = n;code = 110 char = d;code = 100 char = ;code = 32 char = p;code = 112 char = r;code = 114 char = e;code = 101 char = v;code = 118 char = e;code = 101 char = n;code = 110 char = t;code = 116 char = ;code = 32 char = s;code = 115 char = t;code = 116 char = r;code = 114 char = u;code = 117 char = c;code = 99 char = t;code = 116 char = u;code = 117 char = r;code = 114 char = a;code = 97 char = l;code = 108 char = ;code = 32 char = a;code = 97 char = l;code = 108 char = i;code = 105 char = g;code = 103 char = n;code = 110 char = m;code = 109 char = e;code = 101 char = n;code = 110 char = t;code = 116 char = ;code = 32 char = o;code = 111 char = f;code = 102 char = ;code = 32 char = s;code = 115 char = o;code = 111 char = m;code = 109 char = e;code = 101 char = ;code = 32 char = p;code = 112 char = o;code = 111 char = r;code = 114 char = t;code = 116 char = i;code = 105 char = o;code = 111 char = n;code = 110 char = s;code = 115 char = ;code = 32 char = o;code = 111 char = f;code = 102 char = ;code = 32 char = t;code = 116 char = h;code = 104 char = e;code = 101 char = ;code = 32 char = r;code = 114 char = R;code = 82 char = N;code = 78 char = A;code = 65 char = s;code = 115 char = ,;code = 44 char = ;code = 32 char = i;code = 105 char = n;code = 110 char = c;code = 99 char = l;code = 108 char = u;code = 117 char = d;code = 100 char = i;code = 105 char = n;code = 110 char = g;code = 103 char = ;code = 32 char = t;code = 116 char = h;code = 104 char = e;code = 101 char = ;code = 32 char = L;code = 76 char = 1;code = 49 char = ;code = 32 char = s;code = 115 char = t;code = 116 char = a;code = 97 char = l;code = 108 char = k;code = 107 char = .;code = 46 char = <;code = 60 char = b;code = 98 char = r;code = 114 char = /;code = 47 char = >;code = 62 result sql = We describe a method to establish chronologies of ancient ribosomal evolution. The method uses structure-based and sequence-based comparison of the large subunits (LSUs) of Haloarcula marismortui and Thermus thermophilus. These are the highest resolution ribosome structures available and represent disparate regions of the evolutionary tree. We have sectioned the superimposed LSUs into concentric shells, like an onion, using the site of peptidyl transfer as the origin (the PT-origin). This spherical approximation combined with a shell-by-shell comparison captures significant information along the evolutionary time line revealing, for example, that sequence and conformational similarity of the 23S rRNAs are greatest near the PT-origin and diverge smoothly with distance from it. The results suggest that the conformation and interactions of both RNA and protein can be described as changing, in an observable manner, over evolutionary time. The tendency of macromolecules to assume regular secondary structural elements such as A-form helices with Watson-Crick base pairs (RNA) and alpha-helices and beta-sheets (protein) is low at early time points but increases as time progresses. The conformations of ribosomal protein components near the PT-origin suggest that they may be molecular fossils of the peptide ancestors of ribosomal proteins. Their abbreviated length may have proscribed formation of secondary structure, which is indeed nearly absent from the region of the LSU nearest the PT-origin. Formation and evolution of the early PT center may have involved Mg(2+)-mediated assembly of at least partially single-stranded RNA oligomers or polymers. As one moves from center to periphery, proteins appear to replace magnesium ions. The LSU is known to have undergone large-scale conformation changes upon assembly. The T. thermophilus LSU analyzed here is part of a fully assembled ribosome, whereas the H. marismortui LSU analyzed here is dissociated from other ribosomal components. Large-scale conformational differences in the 23S rRNAs are evident from superimposition and prevent structural alignment of some portions of the rRNAs, including the L1 stalk.
Publication: Mol. Biol. Evol. 2009 Nov ;26(11)::2415-25. :doi: 10.1093/molbev/msp163.
Publication Date: 2009 Nov
Study Author(s): Hsiao, Chiaolong;Mohan, Srividya;Kalahar, Benson K;Williams, Loren Dean;
Institution: School of Chemistry and Biochemistry, Parker H. Petit Institute of Bioengineering and Bioscience, Georgia Institute of Technology, Atlanta, GA, USA.
Shortcut link to this study:
We describe a method to establish chronologies of ancient ribosomal evolution. The method uses structure-based and sequence-based comparison of the large subunits (LSUs) of Haloarcula marismortui and Thermus thermophilus. These are the highest resolution ribosome structures available and represent disparate regions of the evolutionary tree. We have sectioned the superimposed LSUs into concentric shells, like an onion, using the site of peptidyl transfer as the origin (the PT-origin). This spherical approximation combined with a shell-by-shell comparison captures significant information along the evolutionary time line revealing, for example, that sequence and conformational similarity of the 23S rRNAs are greatest near the PT-origin and diverge smoothly with distance from it. The results suggest that the conformation and interactions of both RNA and protein can be described as changing, in an observable manner, over evolutionary time. The tendency of macromolecules to assume regular secondary structural elements such as A-form helices with Watson-Crick base pairs (RNA) and alpha-helices and beta-sheets (protein) is low at early time points but increases as time progresses. The conformations of ribosomal protein components near the PT-origin suggest that they may be molecular fossils of the peptide ancestors of ribosomal proteins. Their abbreviated length may have proscribed formation of secondary structure, which is indeed nearly absent from the region of the LSU nearest the PT-origin. Formation and evolution of the early PT center may have involved Mg(2+)-mediated assembly of at least partially single-stranded RNA oligomers or polymers. As one moves from center to periphery, proteins appear to replace magnesium ions. The LSU is known to have undergone large-scale conformation changes upon assembly. The T. thermophilus LSU analyzed here is part of a fully assembled ribosome, whereas the H. marismortui LSU analyzed here is dissociated from other ribosomal components. Large-scale conformational differences in the 23S rRNAs are evident from superimposition and prevent structural alignment of some portions of the rRNAs, including the L1 stalk.
PMID: 19628620